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Tht Tight iif XnatMtation U rttmd. 




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KoDBD THB WoKlp. Wllh IM IlluBliatloDS by PniTCHETT. 2ii. Poptilar 


TION i or. The Pbimksvatioh of Favodki^o Racbs m thb SrKuoc.LK i-OR 

Life. Lirge Tjps EdIUoo, wilh Ponrstt. i vols. lEi. Popular Sdilion, ei. 


FERTILIZED BY ISSlXJra, WoodcaH. U. ed. 


ilESilCATIOH. njuBHntiona. 15s. 

- SES. lUustntlons. Larga Typa Edition, 3 vola. isi. Foimlar Sdition, 

' Vl. M. 

ANIMALS, lllnstmllons. 12<. 
INSECTIVOROUS PLANTS, lllustratious. 9>. 



MONOGRAPH OF THE CIIiRIPEDIA. lUastrationg. 2 vols. 




DoBiNG Ube succoseive reprints of the first edition of this work, 
publiahed in 1871, 1 was able to introduce several important 
corrections; and now that more time has elapsed, I h&-t» 
cndeaTonred to profit by the fiery ordeal through which. Uie 
book has passed, and have taken advantage of all the criticisms 
whicli seem to me sound. I am also greatly indebted to' a Ifitge 
number of eorrespondenta for the communication of a snipriSing 
number of new facts and remarks. These have been so Dnmerone, " 
that I have been able to use only the more important ones ; and 
of these, as well as of the more important corrections, I will 
append a list Some new illustrations have been introdnced, 
Mid foui' of the old drawir^ have been replaced by better ones, 
done from life by Mr. T. .W. Wood. I roust especially call 
attention to some observations which I owe to the Mndnees of 
Prof. Hnxley (given as a suppleraent at the end of Part I.), on 
the nature of the differences between the brains of man and the 
higher apes. I have been particularly glad to give these obser- 
vations, because during the last few years several memoirs on the 
Bubjeot have appeared on the Continent, and their importance 
has been, in some cases, greatly exaggerated by popular writers. 
I may take this opportunity of remarting that my critics 
frequently assume that I attribute all changes of corporeal 
structure and mental power exclusively fo the natural selection 
of such variations as are often called spontaneous; whereas, 
even in the first edition of the 'Origin of Species,' I distinctly 
stated that great weight roust be attributed to the inherited 
efifeets of nse and disnse, with respect both to the body and 
mind. I also ottribnted some amount of modification to the 
direct and prolonged action of changed conditions of life. Some 
allowance, ton, must be made for occasional reversionB ol 


vi Preface to the Second Edition. 

Btniotuie; nor must we forget what I have called '■'eoiTDiateii" 
grotrth, meaning, thereby, that -various parte of tho otganisation 
are in some unknown manner so connected, tliat when one part 
varies, so do others ■ and if voriationa in tho one are accn- 
malaied by selection, other parts will be modified. Again, it 
has been said by several critics, that when I found that many 
details of Btructure in man could not be explained through 
natural selection, I invented sesnal selection ; I gave, however, 
a tolerably dear sketch of this principle in the first edition ol 
tho ' Origin of Species,' and I there stated that it was applicable 
to man. This Eubject of sexual selection haa been treated at 
full length in the present work, simply because an opportunity was 
here first afforded me. I have been struck with tho likeness oi 
many of the half-favourable criticisms on sexual selection, with 
those which appeared at first on natural selection; nuch as, 
that it would explain some few detoils, but certainly was not 
applicable to the extent to which I have employed it. My 
conviction of the power of sexual selection remains unshaJcen ; 
but it is probable, or almost certain, that several of my con- 
clusions will hereafter be found erroneous; this can hardly fail 
to be the case in tho first treatment of a subject. When 
naturalists have become familiar with the idea of sexual selection, 
it will, s& I believe, be much more lai^ly accepted; and it 
haa already been folly and favourably received by eeveral capable 












\ hnman ear reviaed. 



Cases of men born with hairy bodies. 

27. note. 

20, note. 

Mante|:;aZEa oa the laat molar tooth in uao. 



82, note. 

21, note. 

j plftlned by adaptalioa on medmoical 
Intelligence in a bnbooa. 





Sense of hnmour in dogs. 



(FnrUier fiicia on imitation in man and 





Aoqniaition of experiencB bj animala. 





/Power of fotmiDg concepts in relation to 
fPIeasuie faaa certain Bounda. eoloim. and 


{ forms. 



FideUty in the olopiiant. 





Parental afiecUon. 



\ note. / 

Peisifltence of enmity and hatred. 



jKstnre and Mrength of shame, ««Tet, tad 


117, note. 

Suicide amongst saragea. 


120. note. 



Belection, as applied to primeTal man. 
Besemblauces Iwtween idiots and animaZ& 



124, note. 

39. note. 

Division of the malar bone. 


36-8, note. 



Further tasce of muscles proper to animals 

Brooft : average capacity of sknll diminlshea 






TabU of 

he Principal Additions and 











/Disappeaiance of the toil in man and ceiiain 
\ monioys. 



flnjimoHfl forma of eeleotion io oivilised 



/Indolence of man, when free from a n'ao.eAa 



(Gorilla protecting himaalf from rain with hia 

208. note. 

161, note. 





(Changed conditions lessen fertility and cause 
\ iU-bealtb amongst sftTagea. 





(Note by Professor Haxley on the derelop- 
\ ment of the brain in mm trad apes. 



/Special organs of male paiMBitio worms for 
\ LoMing the female. 


jGrcatec vaiiabiHtT of male than femaJo; 




on birds' heads determines their tnins- 
mission to one or both seiea. 



Causes of eiceas of male births. 



Pcoportioii of the seiea in the bee familT. 
(Eicess of males perhaps sometimes detei- 



\ mined by selection. 



Blight colours of lowly organised animals. 



SeiuftI selection amongst spidera. 











Musical appiiiatus of Homoptera. 




288, nolo. 

i Orthoplera. *■ ^^^ 



fHermaQC MiUler on sesual differences o( 



Souuds prodaced by moths. 



Display of beauty l)y butterfliea. 


Temale butterflies, tating the more active 
\ pattmconrt8hip,brigbt*Tthanthdrmales. 



Further oases of mimicry in battorflies and 



'Cituse of bright and diversified oolonrs ol 


Corrections to the Present Edition. 

Brush-like ecalee of male Mallotns. 
(Further (acta on oourtehip of fishes, and the 
I spawning of Mw^ropu^ 

DufoBB^ on the sounds mode by fishes. 

Belt OQ B frof; protected by bright colouring. 

Ifurther facta on mental powers of snakes. 

Sounds produced by snakea; the rattlesnake. 

Combats of Chameleons. 

Marshall on protub trances on birds' hoada. 
fFucthet facta on display by the Argus 
[ pheasant 

Attachment hetircen paired birds. 

Female pigeon rejecting certain males. 

(Albino, birds not finding partners, in a state 
of nature 
Direct action of olimalo on birds' colours. 
/Fm-ther &cts on the ocelli in the Argus 
\ pheasant. 

ITaste foe the beautiful permament enough 
to allow of aexual selection with the Inner 
I animals. 

(Horns of Bheep or^inaliy a masculine 
\ character. 

Cfistration affecting boms of animals. 

Prong-bomed variety of Cervut rirginianat. 

ie and female whales ai 

(Belatiye ai 

Absenca of tuaka in male miocene pigs. 

Dcbsou on sexual differences of bats. 

Eeeks on advantage from peculiar colouring. 
fDiffeience of complexion in men and nomeQ 
[ of an African tnbe. 

Speech aubseqnent Co singing. 

(Schopenhauer on importance of courtship to 

{ mankind. 

fEevision of difcusaion on communal marriagea 

{ and promiscuity. 

(Power of choice of woman in marriage, 

I amongst savages. 

[LoDg-contiuued habit of plucking out haira 

[ may produoe an inberitod effect. 





Teib ErmmicB or thb Descent oy Mah rsOM bohe Lotteb Fobm 

Nature of the evidence bearing on the origin of man — Homologotta 
atractares in man and the lower animals — Mi3f«llaneoua poinla 
of eorreapondenoe — Development — Budimentary strnctureB, 
muBclefl, sense-organs, hair, bonea, reprodnctive organs, fc. — 
The bearing of theee three great olsaseB of facta on the origin of 


O.N THB Manner of Dbtblopment of Man reoa bomb Loweb 

Variability of body and mind in man— Inheritance — Causes of 
variability — Laws of variation the same in man as in tlio iow« 
animalB— Direct action of the eooditions of life— Effeota of the 
increased use and disuse of parta — Arrested development— Ee- 
veraion — Correlated variation — Kate of Increcse — Checks to 
incieBse — Natural aelection— Man the most dominant animal in 
tlie world— Imporlauco of his corporeal stractura — The cauaos 
which have led to hla becoming erect— Coaseqaent changca of 
Btrocturft— Decrease in size of the canine teeth — Increased size 
■nil altered shape of the skuU — Nakedneaa — Absence of a tail — 
Defenoeleaa eondition of maa S< 



The difference in mental power between the highest ape and the 
lowcat savage, imraense— Cerlain inatinofs in common— The 
emotions — Cnrioaity — Imitfction — Attention — Memory — Imagi- 
nation— Reae(m—Pn)g»eeMva imiaswanient— Tools and weapons 


used bf auimals — Abstntotion, Belf-consciousness — laDgnsge 
— Sense of beantj — Sdief in God, spiiitual ngenoiea, euper- 


Tlie moral senae — Fundamental proposition — The rjualitios of 
social animals — Origin of sooiability— Struggle between opposed 
iustinDts — Han a social animal — Tbe more enduring social in- 
stincts conquer other leas persistent instincts — Tlie social virtnes 
elooe regaided by aaTagea — The ielF-regftrding Tirluea acquired 
at a l»t«r stage of development — The importdnce of ihe judg- 
ment of the Eoembers of the same oommnaily on conduct- 
Trail smission of moral tendonoies— Summary 


Advancement of the intellectual pow«rs through natnral selec- 
tion — Imporianca of imitation — Social and moral faoolties— 
Their development within the limits of the same tribe — Natural 
Eelection>as afleoting civilised nations — Evidence that civilisL-d 
nations were once barbarous . • . . . 1 


Ok the Affdjitieb and Genealogy oe Man. 

Position of man in the animal series — The natural Ejetem genea- 
loj^ical — Adaptive characters of alight valuo— yarious small 
points of reaemhlance between man and the Quadrumana — 
Runk of moH in the natural ejistem — Birthplace and antiquity 
of man — Absence of fossil connecting-linkH— -Lower stages in 
the genealogy of man, as inferred, firstly from his affinitjea and 
secondly from hia structure — Early andiogynons condition of 
the Vertebrata — Concluaion I 


On the EiCEg oE Han. 

tbe nature and value of specific chaructera — AppUcafion to the 

races of man — Arguments in fevonr of, and opposed to, ranking 

the Bo-callcd races of man as distinct species — BulMrpeoiea— 

MonogeniBls and polygealita — Conreigenae of oharscteT-- 


Nmneraua poinls of resemblance in body and mind betwceu tlia 
most distinct races of man— The state of man when he first 
ipread ovm the earth— Each raoe not descended &om a. eingla 
pair — The extinction of raoea — Tha formation of laocfl — The 
efiects of oroaaing— Slight influence of the direct Hction of the 
condiCioDB of life — Slight or no infioence of natural suleBtioi> 
Sesual selection 1 




BcoolMlary Boximl charactera — Seinal selection — Manner of action 
—-Excess of males — Polygamy — The male alone generally 
modified through sexual selection — Eagerness of the male — 
Variability of the male — Choice exerted by the female — Sexual 
compared with ntitiita! selection — Inheritance at corresponding 
periods of life, at corresponding seasons of the year, and as 
1 imited by sex — Kelations between the seyeral forma of inheri- 
tanoe— -Cansofl why one sex and the joung are not modified 
through sexual selection — Supplement on the proportional num- 
bers of the two sexes throughout the animal kingdom — The 
proportion of the sexes in relation to natural selection . 2' 


These charaoterB absent in the lowest classes — Brilliant colours — 
MoUusca— Annehda — Crustacea, secondary aeiual characters 
strongly deieloped; dimorphism; colour; characters not ai'- 
qniiud beforo matniity — Spiders, sexual colours of ; stridulation 
by the moles — Myriapoda 21 



Ijiversifled stmatureB poaseSBed by the males for sefcsiog Ihe 
femalea— Differences between the aeses, of which the mean- 
ing is not nnderstood— DiffBreooe in size betmeBn the sexes — 
Thjsanuia — Diplera-— Hemiptera— Homoptera, musical powers 


poaaaned by tho maiet ftlone— Orthoptem, nndBal iwdnmenti 
ofthenialeB, nmuhdiveniflBdin etrnetnTo; pugnacl^iooloiin — 
Nonropten texttal difler«ncea In eolom— Hjmeiuipten, |ing- 
nacitj and oolotirs — Coleopten, colonn; fninuhed »ith greftt 
borDB, appBiently ea an ornament ; bafflae ; BtridalatUg oi^uih 
gOQeisUy oommon to both ses«e .... Z 

Itmoi% eoMntied. — Ohoeb liKpaiofrtai. 

Ouuitship of bnttecOieB-xBattlcs— TiofciDg nolaa— Colonra common 
to both Eoxea, or mtae brUliBot in the male« — Examples — Not 
duo to the direot action of the conditions of life — Colomv 

adapted tor protection — Ooloma of motlu — Display — P-ercoptiTe 
powerB of the Lepldoptera — Varlabilitj — CansoB of the diSerence 
in oolooc between the males and females — Mimicry, ffmate 
batterBie* moce brilliantly colonied than the males — Bright 
colonra of caterpillars — Sumraary and oonolnding remarks on 
the seoondary wmal chanioters of inseda — Birds and insects 
compared ....... 3 


Fishes 1 Cotirtehip and battles of the toales— Larger size of the 
females — Halee, bright cdonts and oratunental appen<)ages; 
other strange chatscters — Colours and appendages acquired by 
the males during the breeding;-seaaan alone — Bishes with both 
Boies brilliantly coloured — Protective colours — The less con- 
spicuous eolouis of the female cannot be accounted for on the 
principle of protection — Malo fishes building nesta, and taking 
chai^ of the ova and young. AjiPHiBiAKa: Differences in 
structure and colour betneen the sexes — Vocal organs. Bbp- 
ules; Cheltmians— Orooodilea — Bnakes, coloors in some cases 
proteoUve — Lizards, battlea of — Omameotsl appendages — 
Strange differences in atraotore between the sexes — Golonrs — 
Sexual differences almost aa great as with birds . , S 


Seoomdabt Sexual Ohabaotbrs op Bibds. 

Besual diSerencea — Law of battle — Special weapons — Vocal 

organa— Instrumental mn^o— Love-antics and dances — Deoo- 

ratiouB, permanoit and seasonal — Double and single annual 

iBoi)lt>~-Displ^of rnnamentsbythemslee . . S 



lilioioo exerted bjr tbe female — Lei^IIi of cunrtehip — tJupsbed 
biida — Mental qniditi«s end tasta for the betntlfiil-— Ptefereace 
or antipathy ■hewn hf the female foi paiticulaT mftlea — Veii- 
■bUItjr ofblide — V«ri«tion« gometimOB abrupt — Lawa of Tsria- 
tion — Ifomiatioa of ocelli — GndatioDB of chaiactei — Case of 
Pmwoc^ Aigna phessant, and Utoeticte . . , 4i 


Id to Thy the male* aloiie of some speclea, and both 
eesea of others are brightly onlonred — On aexiiBlIy-liinited 
inheiitance, aa applied to various attnotorea and to brightly- 
coloored plamage — Nidiflcation in relation to colaor — Lose of 
nnptial ptninage doting the wioter .... i^ 


The immature plomage in relation to the obatacter of the plmnage 
in both BCzes when adult — Six claswe of cases — Sexoal differ- 
enoeB between tho malea of olo«ely-aIlied or representative 
speoiea — The female assuming the characters of the mala — 
Flunu^e of the young in rektioQ to tbe aummer and winter 
plumage of the adults — On tbe increase of beauty in the birds 
of the world — Proteotive colonrfng — Oonspiouously-coloured 
birds— Norelty appredatod — Summary of ths fonr chapterB on 
hhd» 41 


Secohdabt Bkidii. OfusaoTEM or SLunuLs. 

Tbe lavr of battl»— Spedal weapons, oooflned to tbe males — Cause 

of absenoe of weapons in the female — Weapons common to both 

sexet, yet primarily acquired by the male— Other uses of snch 

weapons — Tlieir high importenoe — Gre«ter size of tbe male 

Means of defence — Ou tbe preference shewn by either sex in the 
pairing of qnadmpeda Si 




L'ase of the female being more otnamented tlmu Uie meie — 
Colout end orDnmenta dua to ■esnal leleotion — Cfdoor aoqnirod 
Tor the Bake of pniteotlda — Colonr, thougli oommon to both 
sexes, often dne to seinal mleclloD — On the duappeantnce of ' 
■pots and stripes in adult qnadrnpeds — OiithecoloaTBandDniH- 
menti of the Qnadrnmaika — Btnunutty .... Si 

PART in. 


Skoondabt SainAii Chabactebs 03t Ham 
DifTeieiuies betiresn nun uid woman — Cames of sncli differences, 
and of certain charaDtera cammon to both seiet — Law of battle 
— DiSereocea io mental powers, and voice — On the influence 
of beaut; in determioing the marriages of msuklDd — Attention 
paid b; savages to ornaments— Their ideaa of beaut; in woman 
— The tendenqy to exaggerate each natural pecoliaiity . S. 


Sboondabt Sbxdal CBAjtAoms cr Uan — amUmied. 

On the effects of the continned seleetioa of women according to a 
lUiEtitent standard of beaaty io each race — On the causes which 
interfere vith seinal selection tn civilised and savage nations 
— ConditiotiB faroncabia to semal selection during pilmevol 
times — On the manner of action of eeznal selection with man- 
kind — On the women in savage tribes having some power to 
ohooae their husbands — Absence of hair on the bod;, and 
development of the beard — Oolout of the skin — Sanunary . 5 

General Sciiuabc abd CoNOLtrston. 
Uun oonclasioQ that man Is descended fmm some lower form- 
Manner of development — Genealcgy of man — Intelleetnal and 
morel facnltloa — Sexnal selection — Conchiding remarks . 6i 


Hosted by 





Tus uatnio of the following work will be best nnderstood t^ & 
brief Moonnt o£ how it came to be written. During many Teara 
I collected notes <m the origin oi descent of man, without any 
tatcntbn of publishiDg on the subject, but rather with the 
determination not to publish, as I thought that I shonld thoa 
only add to the pKtjudices against my Tiewe. It seemed to me 
Bttffluient to indicate, in the first edition of my ' Origin of 
Speoies,' that by thia work "light would be thrown on the 
" origin of man and his history ;" and thia implies that man must 
be included with other organio beings in an; general conclusion 
respecting hia manner of appearance on thia earth. Now the 
case wears a, wholly different aspect When a uatoralist like 
Carl Togt ventorea to say in his address aa President of the 
National Institution of Geneva (1869), "personue, en Europe 
" su moius, u'ose plus soutenir la creation independanta et de 
" toutes pieces, des eapdcea," it is mamfest that at least a large 
number of naturalists must admit that species are the modified 
descendantB of other species; and this especially holds good with 
the joimgor and rising iiaturaliBts. The greater number accept 
tho.agoncy of natural stloction ; tboujjh some urge, whether with 
justice the future must decide, that I have greatly overrated its 
importance. Of the older and honoured chiefe in natural science, 
many unfortunately are still opposed to evolution in every 

B now adopted by most naturalists, 
I in every other eaae, be followed by 



otheiB who aro not scientific, I have beea led to pat t^^elhoi 
my notes, so as to see how far the general cosclnsions arrived at 
in mj former worfcs were applicable to man. This seemed all 
the more desirable, as I had neTer deliberately applied these 
views to a species taken singly. When we confine our attention 
I to any one form, we are deprived of the weighty arguments 
\ ' derived from the nature of the affinities which connect together 
whole groups of organisms — their geographical distribution iu 
past and present times, and their geological succeesion. The 
homologicai structure, erabrjological development, and rudi- 
mentary organs of a species remain to be considered, whether it 
be man or any oilier animal, to which our attention may be 
directed ; but these great classes of facts afford, as it appears to 
me, ample and conclusive evidence in favour of the principle ol 
gradual evolution. The strong support derived from the other 
arguments should, however, always be kept before the mind. 

The sole object of this work is to consider, firstly, whether 
man, like every other spooies, is descended from some pre- 
eiistlng form; secondly, the manner of his development; and 
thirdly, the value of the difierencea between the BO-called races 
of man. As I shall confine myself to these points, it will not be 
necessary' to describe iu detail the differences between the several 
races— an enormous subject which has been fully discussed in 
many valuable works. The high antiquity of man has recently 
been demonstrated by the lalwuvs of a host of eminent men, 
beginning with M. Boucher de Perthes; and this is the indis- 
ponsablo basis fur understanding his origin, I shall, therefore, 
t4iko this conclusion for granted, and may refer my readers to 
tlio admirable treatises of Sir Charles Lyell, Sir John Lubbock, 
and others. Nor shall I have occasion to do more than to allude 
to the amount of difference between man and the anthropomor- 
phous apes ; for Prof Huiley, in the opinion of most competent 
judfies, has conclusively shewn that in every visible character 
man diffeis less from the higher apes, than these do from the 
lower members of the same order of Primates. 

This work contains hardly any original facts in regard to man j 

but as the couclusioss at which I arrived, after drawing up a 

rough draft, appeared to me interesting, I thought that tiiey 

might interest others. It has often and confidently been asserted, 

^ that man's origin can never be known : but ignomnco more 

f" 'sfe. frequently begets confidence than does knowledge: it is those 

■V ivho know little, and not those who know much, who so 

vely assert that this or that problem will never he solved 

\}>y scienca The conclusion that man is the co-descendant with 

other epecios of some andent, lower, and estinct fonn, is not in 


fytt oduction. 

■ny degree now. Laaiarck long ago came to this conolnsion, 
which has lately bee& maintained by sereiftl erDinent naturalisls 
and philcBophers ; for instance, by Wallace, Hnsley, Lyell, Togt. 
Lubbock, Biichaei, Rolle, &c.,' and especially by Hackei. This 
last naturalist, besidea his great work, 'Generelle Morpholo^e' 
(1866), has recently (1868, with a second edit, in 1870), pub- 
lished his ' Naturliche Schijpfungseeschichte,' in which he fully 
diseuaaea the geneali^y of man. If this work had appeared 
before my essay had been written, I should probably never have 
completai it. Almost all the conclusions at which I have 
arrived I find confirmed by this naturalist, whose knowledge on 
many points ia much fuller than mine. Wherevei I have added 
any fact or view from Prof. Hackel's writicgs, I give his autho- 
rity in the text ; other statements I leave as they originally stood 
in my manuscript, occaaionaO j giving in the foot-notes references 
to his works, aa a confinnation of the more doubtful or interestii^ 

During many years it has seemed to me highly probable that 
sesual selection has played an important part in differentiating 
the races of man ; but in my ' Origin of Species ' (first edition, p. 
199) I contented myself by merely alluding to this belief. When 
I came to apply this view to man, I found it indispensable to 
treat the whole subject in full detail.' Consequently the aecond 
part of the present work, treating of sesual selection, has ex- 
tended (o an inordinate length, compared with the first part; 
but this could not be avoided. 

I had intended adding to the present volumes an essay on the 
expreasion of the various emotions by man and the lower animals. 
My attention was called to this subject many years ago by 
Sir Charles BcU'b admirable work. This illustrious anatomist 

■ned Nat.,' Modena, 16S7, p. 81) a very 

not girs the titles ; bat aa those 

GugUnd, I will gire them :—' Scul 
VoclesUBgen tlber die Darwin' ' 

Theorie :' zweite Au9age, 18B8, tod 

of " Man, made in the image of i 

Dr. L. BUohneri translated into 

" waa also made in tlie image ot 

French nnder the title 'Confa-eDCPS 

' Prof. Hackei was the ■ 

'Der Menach, im Lichte der Dal- 

author who, at the time when 

win'schB Lehre,' ISHS, vod Dr. F. 

work first appeared, had discu 

Euile. 1 will not attempt to ^ive 

the subject of sexual selection, 

had seen its full importance, c 

hara taken the Bsme side of tha 

the pabllcation of the ' Origin ■; 

question. This G. CaDestrini has 

this he did in a very able mann. 

published ('Ann-ario delta Soc. X. 

kii vuions vork& 


4 Introdtiction, 

maintains that man is endowed with certain mnsclea solely for 
tho Bake of expressing his emotions. As this view is obTionsIy 
opposed to the belief that man is descended from some other and 
Jower form, it was necessary for me to consider it I litewise 
wished to ascertain how far the emotions aro cspreased in the 
same manner by the different races of man. But owing to the 
length of the present work, I hare thought it better to TC&ane 
mf essay for separate publicitioii. 


( 5 ) 

Part I. 

Sataie ot the evidence beariDg on th« origin of mao — Horaologoni 
■tructuies in Biau sad the lower animals — Miacellaneons poinu ol 

organs, bail, bonea, repiodnctire orgaoE, &c — Tbe bearing of these three 
great classes of ^ts oa the origin of man. 

He who wishes to decide whether man is the modified descendant 
of some pre-esisting form, would prabahly first eBquire whether 
man varies, howeyer eUghtly, ia bodily Etructuro and in mental 
fiiCTiItiee; and if so, whether the vaiiatioas are transmitted to 
his offspring in oooordanee with the laws which preTail with the 
lower animals. Again, are the variations the result, as far a^ 
onr ignorance pennita us to judge, of the same general causes, 
and are they governed by the same general laws, aa in the case 
of other organisms; for instance, by correlation, the inherited 
effects of use and disuse, &c. ? Is man subject to similar mal- 
conformations, the result of arresicd development, of reduphcation 
of parts, &o., and does he display in any of his anomalies rever- 
sion to some former and ancient type of structure ? It m^ht 
also naturally be enquired whether man, like so many other 
animals, has given rise to varieties and sub-races, differing but 
elightly from each other, or to races differing so mnch that fltey 
mnst be classed as doubiflil species? How are saeh races 
distributed over the world; and how, when crossed, do they 
react on each other in the first and sncceeding generations ? [ 
And so with many other points. 

The enquirer would nest come to the important point, 
whether man tends to incicaee at so rapid a rate, as to lead to 
occasional severe atrnggles for existence; and consequently t« 


6 The Descent of Man. Part !, 

beneficial Tsriations, whether in body or miad, being preserved, 
and injurious ones eliminated. Do tlie races or species of men, 
whichever term may be applied, encroach on and replace one 
another, so that some finally become extinct ? We shall eee that 
all these questions, as indeed is obvious in respect to most of 
them, must be answered in the afSrmative, in the same manner 
as with the lower animals. But the several considerations just 
referred to may be conveniently deferred for a time ; and we 
will first see how far the bodily structuro of man shows traces, 
more or le^ plain, of his descent from some lower form. In ,' 
Hucceeding chapters the mental powers of man, in comparison) 
with those of the lower animals, will be considered. | 

Thi Boiily Structure of Man. — It is notorious that man is 
constructed on the same general type or model as other mnm- 
mals. All the bones in his skeleton can be compared with 
corresponding bones in a monkey, bat, or seal. 80 it is with his 
muscles, nerves, blood-vessels and internal viscera. The braio, 
the most important of all the oi^ns, follows the same law, as 
shewn by Huxley and other anatomists. Bischoff,' who is a 
hostile witness, admits that every chief fissure and fold in the 
brain of mau has its analogy in that of the orang ; bnt he adds 
that at no period of development do their brains perfectly agree ; 
nor could perfect ^reement be expected, for otherwise their 
mental powers would have been the same. Vulpian ' romaxkH : 
" Lea differences reelles qui esiet«nt entre I'encephale de 
" ITiomme et celui des singes superieurs, sont bien minimes. II 
" ue &ut pas se faire d'illusions k cet Sgard. L'hommo est bien 
" plus prSs des singes anthropomorphes par Ics caracteres 
" anatomiques da eon cerveau que cem-ci ne le sont non- 
"seulement des autres mammiferes, raais m&ne de certains 
" qnadrumanes, dea guenons et des macaques." Bnt it would 
be superfluous here to give further details on the corrospondeDce 
between man wid the higher mammals in the structure of the 
brain and all other parte of the tiody. 

It may, however, be worth while to specify a few points, noi 
uirectly or obviously connected with structure, by which this 
correspondence or relationship is well shewn. 

Jlan is liable to receive from the lower animals, and to com- 

1 -GrosshiiniMQdungen des M^u- 

in t],e Preface to this edition. 

Mhen.' 1868, s S6 The conclusion. 

■ 'U?.sorlaPrijs.'J866,p.890, 

of this author, as well as th >se of 

as quoted by M. Dally, ' L'Orire des 

Primates et le TranaformiamB," IMS, 

brain, will fcn discuMod bj Prof 

p 29. 

Hoilev in the Apfenilij aJludisl to 


Cha?. I. 

Hontological Structures, 

municaie to them, certaiu diseases, as hydToptobia, Tariola, tlie 
glanders, ejpliilis, cholera, herpes, &c. ; ^ and this fact proves thu 
close Bimilarity* of tfieir tissues and blood, both in minuto 
structuxo and composition, far more plainly than does their 
comparison under the best mieroaeope, or by the aid of the best 
chemical analysis. Monkeys are liable to many of the same non- 
eoatagions diseases as we are; thus Keagger,' who carefully 
observed for a loi^ time the Cehvs Azarce in its native land, 
found it liable to catarrh, with the nsual symptoms, and which,' 
when often recurrent, led to consumption. These monkeys 
suffered also from airaplexy, inflammation of the bowels, and 
eatnract in the eye. The younger ones when shedding their 
milk-teeth often died from fever. Medicines produced the same 
effect on them as on us. Many kinds of monkeys have a strong 
taste for tea, coffee, and spirituouiS Uqnors ; they will also, as I 
have myself seen, smoke tobacco with pleasure." Brehm asserts 
that the natives of north-eastern Africa catch the wild baboons 
by exposing vessels with stroi^ beer, by which they are made 
diunk. He has seen some of these animals, which he kept in 
confinement, in this state ; and tie gives a laughable account of 
their behaviour and strange grimaoes. On the following 
momii^ they were very cross and dismal ; they held their aching 
heads with both hands, and wore a most pitiable expression: 
when beer or wine was offered them, they turned away with 
disgust, bat relished the juice of lemons.' An American moniey, 
an Ateles, after getting drunk on brandy, would never touch it 
again, and thus was wiser than many men. These trifling facts 
prove how similar the nerves of taste must bo in monkeys and 
man, and how similarly their whole nervous system is affected. 
Man is infested widi internal parasites, sometimes causii^ 

■. W. Under Lim 

t fluids by the a 

le chomicfd 

rented this su)>ject at some ieagth 


n tha 'Journal of Mental Science,' 

> ' ;:JatuiEeschichte der Sauge- 

nly 1871; Bud in the 'Edinburgh 

thiere voq Paraguay," 1830, s. 50, 

eterinai'7 Revianr,' July 1B5S. 

• A Reviewer has criticised 

some animals much lower in the 

British Quarterly Keview," Oct. 

Bc:^le. Mr. A. Nicols informs oie 

St, 1871. p. 47i!) what I have here 

that he >ept in Qoeenslaud, in Aus- 

, three iaiividuals of the 

roJarcias cineretu ; nni thai. 


lut having been tanght in any 

w they Hcquiied, a strong t 

■■ '■ 'Thierleben.'B.i. 18IJ4, 
)n the Ateles, s. 105. 
s logons Gtateiuects, s«9 


8 The Descent of Man. Vixi L 

fatal efTectfi; and is plagued by external parasiMS, all of wliicb 
belong to the same genera or families as those infesting other 
mammals, and in the case of scabies to the same species.' Man 
is sulgeet, like other mammals, birds, and even insects,' to that 
mysterious law, which causes certain normal proeesBes, snch as 
gestation, as well as the maturation and duration of rarioua 
diseases, to follow lunar periods. His wounds are repaired by 
the same process of healing; and the stumps left after the 
amputation of his limbs, espeoistOy during an early embryonio 
period, occasionally possess some power of regeneration, as in 
the lowest animals.'" 

The whole process of that most important function, the 
reproduction of the species, is strikingly the Same in all mam- 
mals, from the first act of courtsBip by the male," to the birth 
and nurturing of the young. Monkeys are born in almost as 
helpless a condition as our own infants ; and in certain genera 
the young differ fully as mnch in appearance from the adults, aa 
do our children from their full-grown parents.'^ It has been 
ui^ed by some writers, as an important distinction, tliat with 
man the young arrive at maturity at a much later age than with 
any other animal : but if we look to the races of mankind wtiich 
inhabit tropical countries the difference is not great, for the 
orang is believed not to be adult till tlie age of from ten to fifteen 
years."" Matt differs from woman in size, bodily strength, 
hairiness, &o., as well as in mind, in the same manner as do the 

iriodioity," ' British i 

1842. Dr. MaccaMoch, ' . . 

Korth Americdn Jonroal of Science,' " aihil tnrpius potest ii 

vol. xvli. p. 305, has seen t, iag " omnia hominibns et Quadrumani* 

auSeiing from tertinn ague. Here- " commuDia. Nai'rat enim Cyno- 

aftflr 1 shall return to thta aubjeit. " cephalum qnendain in furorera ia- 

" 1 have givea the evidence on " cidere aspecta feminarum ali- 

this head in' xaj ' Variation of Ani- " quarum, aed Dequaquam Bccendi 

tioa,' vol. !i. p. 15, and nors could " per eligebat Juniores, et dignos- 

be added. " oebat in turbS, et advocaliit voce 

" " Mares e dlversis generibas " gestflque." 

" Quadrumanonim sine dubio di- " This remark is made with ra- 

" gnoscunt feminas huraanas a ma- spect to Cynocephatus and the bd- 

" ribus. Primura, credo, odoratu, tiiropomorphoos BpBS by Geoffmj 

•■ poBWa Bspeeto. Mr. Youatt, qui Salat-Hil^re and F. Oavier, 'Hist. 

" din in Hortis Zoalogicis (Besti- Nat. des llammit^ree,' torn. i. 1824. 


Chap. I, Homological Structures. 9 

two sexes of many mammals. So that the correspondence in 
general structure, in the minute Btructure of the tisauea, ic 
chemical composition and in constitution, hetween man and the 
higher auimals, espeoittlly tho anthropomoiphous apea, is ex- 
tremely close. — ' 

Emlryonic Developmeni. — Man is developed from an ovule, 
about tbe 12Sth of an inch in diameter, which difiera in no 
respect from the ovules of other aajirnds. The embryo itself at 
a very early period can hardly be distinguished from that of 
other memliers of the vertebrate kingdom. At this period tlie 
arteries nm ia arch-like branches, as if to carry the blood to 
hranchite which are not present in the higher vertcbrata, though 
tho slits on the eides of the neek stiO remain (/, g, fig. 1), 
Kiarkiug their fonner position. At a somewhat later period, 
■when the extremities are developed, " the feet of lizards and 
" mammals," as the illustrious Von Baer remarks, " the wings 
" and feet of birds, DO less than the hands andifeet of man, all 
" arise from the same fondamental form." It is, says Prof 
Huxley," " quite in the later Btagcs of development that the 
young human being presents marked differences from the yotuig 
" ape, while the latter departs as much from the d<^ in its 
" developmentB, as the man does. Startling as this last assertion 
" may appear to be, it is demonstrably true." 

As some of my readers may never have seen a drawing of an 
embryo, I have given one of man and another of a dt^, at about 
the same early stage of development, carefully copied from two 
works of undoubted accuracy." 

After the foregoing stat^ents made by such high autho- 
rities, it would be STiperfluous on my part to give a number ot 
borrowed details, shewing that the embryo of man closely 
resembles that of other mammals. It may, however, be added, 
that the human embryo likewise resembles certain low forms 
when adult in various points of structure. For instance, the 
heart at first exists as a simple pulsatii^ vessel; the excreta 
are voided through a cloacal passage ; and the os coccyx projects 

" ' Mflo's Place in Natnre,' 1S63, magnified, the embi-jo being twenty- 

p. 67. five days oil The internal viscera 

" The hnman embryo (apper h3vsbeeaomitted,aiidtheuUiiaeap- 

lig.) is from £cker, ■ Icones Phys.,' pend^ee ia both drawings remoTed. 

18S1-18&9, tab. iii. fig. 2. This I was di/etted to these figoi-ea Ij 

embryo was t*n lines in length, so ''"-' ■"--'— ' — — i... 1 

The embryo of the dog is from 
llischofF, ' Entwicklungageschiclite 
dm Hiinde-Eies,' 1845, tab. si. fig. 
13 B This drawing is live times 


The Descent of Man. Part I, 

Fig. I. Upper %nrf human tn bryn f m lickpr, Lowerfigiire llialof»di& 

re-br^n, cerebral hemiApberdp, &c 



;!n*p. I. Rudimsnts. i\ 

lito a true tail, "estendii^ eonBiderably beyond tho rudi- 
" mentaij I^s."'* In the embryos of all air-breathing verteljrateB, 
certain glands, called the corpora WoifBana, correspond with, 
and act like the kidneys of mature flshos.'^ Even at a later 
embryonic x>eriod, some strikit^ resemblances between man aiul 
the lower animals may be observed. Bischoff says that the 
convolntiona of the brain in a human foetus at the end of the 
seventh month reach about tlie same stage of development as in 
a baboon when adult,'" The great toe, as Prof. Owen remarks," 
" which forms the fulcrum when standing or walking, is 
'* perhaps the most characteristic peeubarity in the human 
structure;" but in an embryo, about aa inch in length, Praf. 
Wyman" foimd"that the great toe was shorter than the others ; 
" and, instead of being parallel to them, projected at an angle 
" from the side of the foot, thus corresponding with the per- 
" manent condition of this part in the quadrumana," 1 vrill 
conclude with a quotation from Eusley,^ who after aakii^, 
does man originate in a different way from a dog, bird, frog or 
fish? says, "the reply is not doubtful for a moment; without 
" question, the mode of origin, and the early stages of tho 
" development of man, are identical with those of the animals 
" immediately below him in the scale : without a doubt in 
" these respects, he is far nearer fo apes than the apes are to 
" the dog." 

Hudimejifs. — This subject, though not intrinsically more 
important than the two last, will for several reasons be treated 
here more fuUy.^ Not one of the higher animals can be named 
which does not bear some part in a rudimentary condition ; and 
man forms no esception to the rule. Eudimentaiy organs must 
be distinguished from those that are nascent ; though in some 
eases the distinction is not easy. The former are either abso- 
lutely useless, such as the mamm» of male quadrupeds, or the 
incisor teeth of ruminants which never cut through the gums ; 
or they are of such slight service to their present possessors, 
that we can hardly suppose that they were developed under tho 

" Praf. Wyman in ' Pi-oe. of " I had written a rough copy of 

Amerkan Acad, of Sciences,' sol. iv. this thapter before reading a valu- 

1860. p. 17. able paper, "Car8ttsi-l rudirDentali 

" Owen, 'Anatomy of Verte- in ordine all" origins del nomo" 

bcates,' vol. i. p. 533. (' Annnario della Soc d. Nat.,' Mo-' 

"'Die Grosshirnwindungen dea dena, 1867, p. 81), by Q. Canestria, 

Ueoechen,* 1S68, s. 95. to which paper 1 am considerably 

'* 'Anatoiay of VeTtetrates,* vol. indebted. Hackel has given admir- 

li. n, 553. ahlB diaeusMons on this whole tub. 

» ' Proc Soc. Nat. Hist." Boston, ject, under the title of DystelBology, 

19S3, ToL ii. p. 185. in his ' Oflnerelle Morphologie ' nud 

-' ' Man'a Place in Nature,' p. 65, ' SchSpftmgFgescliichta.' 


1 2 The Descent of Man. Pinr 1, 

eonditioDS which now exist. Organs in this latter state are not 
Ktriefly rudimentary, but they are tending in this direction. 
Nascent oi^ans, on the other hand, though not fully deteloped, 
aro of high service to their i>0i5sessore, and are capable of further 
development. Eadimentaiy organs are eminently Tariable ; and 
this is partly intelligible, as they are useless, or nearly useless, 
and consequently are no longer subjected to naturaJ selection. 
They often become wholly suppressed. When this occurs, they 
are nevertheless liable to occasional reappearance through 
reversion— a circumstance well worthy of attentiou. 

The chief agents in causing organs to become rndimenlary ■ 
seem to have been disuse at that period of life when the organ 
is chiefly used (and this is generally during maturity), and also 
inheritance at a corresponding period of hfe. The tei-m 
"disuse" does not relate merely to the lessened action of 
muscles, but includes. a diminished flow of blood to a part or 
organ, from being subjected to fewer alternations of pressure, or 
from becoming in any way lees habitually active. Eudimenta, 
however, may occur in one sex of those parts which are normally 
present in the other ses ; and such rudiments, as we shall 
hereafter see, have often originated in a way distinct from those 
hero referred to. In some cases, oi^ans have been reduced by 
means of natural selection, from having become injurious to the 
species under changed habits of life. The process of reduction 
's probably often aided through the two principles of compensa- 
tion and economy of growth ; but the later stages of reduction, 
after disuse has done all that can fairly be attributed fo it, and 
when the saving to be effected by the economy of growth would bo 
very small,^ are difScult to understand. The final and complete 
suppression of a part, already useless and much reduced in size, 
in which case neither compensation nor economy can come into 
play, is perhaps intelligible by the aid of the hypothesis of 
pangenesis. But as the whole subject of rudimentary oi^ns 
has been discussed and illustrated in my former works," I need 
here say no more on this head. 

Rudiments of various muscles have been observed in many 
parts of the human body ;" and not a few muscles, which are 

" Some good criticisms on this Zoolog. 1853, tom. iviii. p. 13) de- 
BubjECt have been giTeo by Messrs. scribes ' " .... 

Mai'ie snit Uivart, in < tranaact. nbat h 

Plants nnder Domestication,' vol. ii. muscle, called " le tibial postSrieur," 

pF.317aad397 See also 'Origin is gonenilly qoite abaent in Ihi 

of Species,' 5th edit. p. 535. hnad, but appears from time to ttox 

" For inslaoce M, Riengrd (' An- in a more or less rudimentary cos 


Ceap. 1. Rudiments. 1 3 

regularly present in some of the lower animals can oecasionallj 
be detected ia man in a greatly reduced condition. Every one 
must have noticed tho power which many ammaJs, espeeiallsw 
horses, possess of moTisg or twit«hing their skin; atid tliis is 
effected by the panrdculus cammus. EemtiantB of this muscle 
in an efficient stafe are found in TarioAis parts of our hodios ; for 
instance, the musfle on the forehead, by wliich the eyebrows are 
raised. The plaiyama myoidea, which is well developed on the 
neck, belongs to this sjstem. Prof. Tamer, of Edinburgh, has 
occasionally detected, as he informs me, muscular fasoicali in 
five different situations, namely in the asiljje, near the acapulra, 
iSc, all of which must be referred to the system of the panni- 
cidus. He has also shewn'" that the musculus siernalis or siemalis 
hruloritm, which is not an extension of the reclus abdominalh, 
but is closely allied to tho panniadus, occurred in the proportion 
of about three per cent, in upwards of 600 bodies : he adds, that 
this muscle affords "an excellent illustration of tho statement 
" that occasional and rudimentary structures are especially 
" liable to variation in arrangement." 

Some few persons have the power of contracting the super- 
ficial muscles on their scalps ; and these muscles are in a 
variable and partially rudimentary condition. M. A. de Candolle 
has communicated to me a curious instance of the long-continued 
persistence or inheritance of this power, as well as of its unusual 
development. He knows a family, in which one member, the 
present head of the family, could, wlien a youth, pitch several 
hea,vj books from his head by the niovement of the scalp alone ; 
and he won wagers by performing this feat. His father, uncle, 
r, and his three childxen possess the same power to 
J unusual degree. This faroOy beenmo divided eight 
4 ago into two branches; so that the head of the 
above-mentipned branch is cousin in the seventh degree to tho 
head of tho other branch. This distant cousin resides in 
another part of Prance ; and on being asked whether he possessed 
the same faculty, immediately exhibited his power. This case ofiers 
a good' illustration how persistent may be the transmission of an 
absolutely useless faculty, probably derived from our remote semi- 
human progenitors; since many monkeys have, and frequently 
use the power, of lai^ly moving their scalj® up and down.^' 

The extriiMic muscles which serve to move tho external ear, 
■md the intrinsic muscles which move the dilferent parts, are in a 
rudimentary condition in man, and they aJl belong to the sysiem 

'■ Prof. W. Turner, ' Proc. Rojal Emotions in Uon and AnimalV 
Soc. Edinburgh,' 1868-67, p. ;5. 1873, p. 144. 

*' Sm my ' Eipreaaioa ' f the 


t4 The Descent of Matt. Paht 1 

of the pnmniouJits ; thej are also variable in duvelopment, or at 
least in function, I have seen one man wlio could draw thy 
whole ear forwards; other men can draw it upwards; another 
who could draw it backwards;^ and from what one of these 
persons told me, it is probable liiat most of us, by often touching 
our ears, and thus directing our attention towards them, could 
recover some power of movement by repeated trials. The power 
of erecting and directing the shell of the ears to the various 
pointa of the compass, is no doubt of the highest service to 
many animals, as thej thus perceive the direction of danger i 
but I have never heard, on sufBeient evidence, of a man who 
possessed this power, the one which might bo of use to him. 
Tlie whole estcrnal shell may be considered a rudiment, tc^etier 
with the various folds and prominences (helis and anti-helix, 
tragus and anti-tragns, &c) which in the lower animals 
strengthen and support the ear when erect, without adding . 
much to its weight. Some authors, however, suppose that the 
cartilage of the shell serves to transmit vibrations to tha 
acoustic nerve; but Mr. Toynbee,™ after collecting aH the 
known evidence on this head, concludes that the external shell 
is of no distinct use. The ears of the chimpanzee and orang aw 
curiously lite those of man, and tie proper muscles aie lifcewiso 
hut very shghtly developed,™ I am also assured by the keepers in 
tho Zoological Gardens that these animals never move or erect 
their ears; so that they are in an equally rudimentary condition 
with those of man, as far as function is concerned. Why these 
animals, as well as the progenitors of man, should have lost tho 
power of erecting their ears, we cannot say. It may be, though 
I am not satisfted with this view, that owing to their arboreal 
habits and great strength they wore but little eiposed to danger, 
and so during a lengthened period moved thoir ears but little, 
and thus gradually lost tho power of movmg them. This 
would be a parallel case with that of those large and heavy 
birds, which, from inhabiting oceanic islands, have not been 
exposed to the attacks of beasts of prey, and have consequently 
lost tiio power of using their wings for flight. The inability to 
move the ears in man and several apes is, however, partly com- 
pensated by tho freedom with which they can move the head in 

1 the 

» CanestriniquoteaHyrtl, ('An- 

lately beeo eipfrimeotiog on 

naario delU Soc. del Naturalist!, 

function of the shell of the 

Modena. 1867, p. 97) to the same 

sad hmi come Co nearly the , 


ooEolusion m that giv™ here. 

» 'The Diseases of the Ear,' by 

" Prof. A. Macalister, 'Al 

J. Toynbee, F,R.S,, I860, p. 12. 

and Mag. of Jiat. History,' vol 

1871, p. 343. 

Pwyer, intbraa me that be had 


OHit. t. 



a horizontal plana, so as to catch sounds from all directions. It 
haa been asserted that the ear of man alone possesses a lobule ; 
but "amdiment of it is found in the gorilla;"^ and, as I hear 
from Prof. Prejer, it is not larelj absent in the negro. 
-The celebrated sculptor, Mr. Woolner, informs me of one little 
peculiarity in the external ear, which he has often observed both 
in men and women, and of which he perceived the full signi- 
flcance. His attention was first called to the subject whilst at 
work on his figure of Puck, to which he had given pointed ears. 
He was thus led to examine the ears of various monkeys, and sub- 
sequently more carefully those of man. The peculiarity consists 
in a little blunt point, projecting from the inwardly folded mai^ii, 
or helix. When present, it is develojjed at lirth, aiid,accoiiiing 
to Prof. Ludwig Sleyer, more frequently ' 
Mr. Wooiner made an exact model of 
the accompanying drawing. (Fig. 2.y 
These points not only project inwards 
towards the centre of the ear, but often 

from directly in front or behind The> 
are variable in size, and some^^hat in 
position, standing either a little higher 
or lower; and they sometimes occur 
on one ear and not on the other They i 
are not confined to mankind, for I ob 
served a case in one of the 'ipider 
monkeys (^Atdes beehehuth^ m our 
Zoological Gardens; and Dr. E. Eay 
Xankestor informs me of another case 
in a chimpanzee in the gardens at a. The pn^ccung pidat 
HamWi^. The helix obvioosly con- 
sists of the extreme margin of the ear folded inwards : and 
this folding appears to be ia some manner connected with the 
whole external ear being permanently pressed backwards. In 
many monkeys, which do not stand high in the order, as baboons 
and some species of maeacus," the upper portion of the ear is 
slightly pointed, and the margin is not at all folded inwards ; 
but if the margin were to bo thus folded, a slight point would 
necessaiiiy project inwards towards the centre, and probably a 
little outwards from the plane of the ear ; and this I believe to 

" Mr. St. Geo 

e Mirar 

rnentary An tomy,' 1873, p. 

" See also soms remarl 

the draniags ui tha card 


LeraHroidea, in Messrs. ainti* ana 
Mivnrt'a ejcellent piiper in 'Tran- 
suet. Zoolog. Sofc' vol. vii. 1869, pp 
G audSa 


1 6 Tlie Descent of Man. Fart I. 

be tlieir origin in many cases. On the other hand. Prof. L. Meyer, 
in an able paper recently piibhEhed," maintains that the whole 
case is one of mere variability ; and that the projections are not 
loai ones, but are due to the internal cartilage on each side <A 
the points not having been fully developed. I am quite ready 
to admit that this is the correct explanation in many instancos, 
as in those figured by Prof. Meyer, in ■which there are several 
minute points, or the whole margin is sinuous. I have myself 
seen, through the kindness of Dr, L. Down, the ear of a micro- 
cephalous idiot, on which there is a projection on the ontsido 
of the helis, and not on the inward folded edge, so that this 
point can have no relation to a former apes of the ear. Never- 
theless in some oases, my original view, tliat the points 
are vestiges of tha tips of formerly erect and pointed ears, 
still seems to ino probable. I think so from the frequency ot 
their occurrence, and from the general correspondence in 
position with that of the tip of a pointed ear. In one case, ot 
which a photograph has been sent me, the projection is so large, 
that supposing, in accordance with Prof Meyer's view, the ear 
to be made perfect by the equal development of the cartilage 
throughont the whole extent of the margin, it would liave 
covered fully one-third of the whole ear. Two cases have been 
communicated lo me, one in North America, and the other in 
England, in which the upper margin is not at all folded inwards, 
but is pointed, so that it closely resembles the pointed ear of an 
ordinary quadruped in outline. In one of these cases, which was 
that of a young child, the father compared the ear with the 
drawing which I have given" of the ear of a monkey, the 
Cynopiihecus niger, and says that their outlines are closely 
Bimilar. If, in those two cases, tha margin had been folded 
inwards in the normal manner, an inward projection must have 
been formed. I may add that in two other cases the outline stili 
remains somewhat pointed, although the margin of the upper 
part of the ear is normally folded inwards— in one of them, 
however, very narrowly. The following woodcut (No. 3) is an 
accurate copy of a photograph of the fcetns of an orang (kindly 
sent me by Dr. NifscheJ, in which it may be seen how different the 
pointed outUne of the ear is at thj" period irom its adnit condition, 
when it bears a close general resemblance to tliat of man. It ia 
evident that the folding over of the tip of such an ear, unless it 
changed greatly during its further development, would give riso 
to a point projecting inwards. On the whole, it still seems to 

»■ UeberdasDarwin'seheSpitzohf, " 'The Expression of the Em* 

Archly fllr Path. Anat. ncd Fhye. tions," p. 136. 


Chap. I. Rudiments. 17 

me provable that the points in question are in some cases, both 
In roan and apes, vestiges of a former condition. 

The nictitating membrane, or third eyelid, with its •wseessory 
muscles and other structures, is especially well developed in 
birds, and ia of much functional importance to them, as it can 
be rapidly drawn across the whole eye-halL It is found in some 
reptiles and ampbibians, and in certain fishes, as in sharks. It 
is fairly well developed in the two lower divisions of the mam- 
malian series, namely, ia the monotremata, and marsupials, and 
m some few of the higher mammals,- as in the wairus. But in 
man, the quadrumana, and most other mammals, it exists, as is 
admitted by all anatomists, as a mere mdiment, called the 
semilunar fold." 

The sense of smell is of the highest imjrartance to the greater 
number of mammals— to some, as the ruminants, in warning 
them of danger; to others, as the carnivora, in finding their 
prey; to others, ;^ain, as the wild boar, for both purposes 
combined. But the sense of smell is of extremely slight service, 
if any, even to the dark coloured races of men, in whom it is 

" Holler's ' Eiemenls of Physi- Knoi, ' Great Artists nnd Anato- 

olojy,' Eng. translat., 1843. vol. 11. mists,' p. 106. This rudiment ap- 

p. HIT. Owen, ' Anatomy of Verte- parently ia somewhat larger in 

bratea,' toL iii. p. 260; ibid, on Negroes and Australians than in 

the Walrus, 'Proo. Zoolog. Soc' Europeans, see Cail Vogt, ' Lectures 

NorembcT 8th, 185*. See also R. on Man,' Eng. translat. p. 129. 


l8 Tke Descent of Man. Paht I. 

much more higiily developed than in the white and ciyilisoj 
races.'* Nevertheless it does not warn them of danger, nor guirfn 
thom to their food; nor does it prevent the Esquimaui from 
sleeping in the most fetid atmosphere, nor many savages from 
eating half-pufrid meat. In Europeans the power differs greatly 
in different indiTldiials, as I am assured by an eminent naturalist 
who possesses this sense highly developed, and who has at- 
tended to the subjects Those who beh'eve in the principle 
of gradual evolution, will not readily admit that the sense of 
smell in its present state was originally acquired hy maji, as 
he now exists. He inherits the power in an enfeebled and 
so far rudimentary condition, from some early progenitor, to 
whom it was highly serviceable, and hy whom it was con- 
tinually used. In those animals which have this sense highly 
developed, such as dogs and horses, the recollection of persons 
and of places is strongly associated with their odour ; and we can 
thus perhaps understand how it is, as Dr. Maudsley has truly 
remarked," that the sense of smell in maa " is sii^ularly effective 
" in recalling vividly the ideas and images of forgotten scenes 
" and places." 

Man differs conspicuously from all the other Primates in being 
almost naked. But a few short straggUng hairs are found over 
the greater part of the body in the maa, and fine down on that 
of the woman. The different races differ much in hairiness ; and 
in the individnals of the same race the hairs are highly variable, 
not only in abundance, but likewise in position : thus in some 
Europeans the shoulders are quite naked, whilst in others they 
bear thick tnfta of hair.'' There can bo little doubt that the 
hairs thus scattered over the body are the rudiments of the 
uniform hairy coat of the lower animals. This view is rendered 
all the more probable, as it is known that fine, short, and pale- 
coloured Lairs on the limbs and other parts of the body, occasion- 

" The occount given by Hnmboldt olfactory region, us well as of the 

t h pow f m 11 p wdby k f the body. I have, therefore, 

h t 63 f tw th Am oa » 11 pok n in the teit of the dark- 

k w d h b hrm d b I red races hating a finer sensa 

h n, M H ( 8 d f mcll thm the white races. See 

les Fic 1 es M tdl s, i t m hi paper,' Medieo-ChimrgicalTran- 

187 p fil) asserts th t h t ns,' Loadon, vol. liii., 1870 

pc dly mad penm U, d p rt. 

p d h ^ gr d IndjiD The'Physiology and Pathology 

Id gn p ns th d k fM nd,' 2nd edit. 1868, p. 134. 

by til od D W Ogl h w Eschricht, Ueber die Richlnne 

"dm b t ns i Haare am measohlichen KSrper, 

th ™ betwee h pow Mflller'a ArchivfUrAnat, und Phys ' 

f m 11 d h 1 g m tte 183 e. 47. I shall often bav* U 

of a m coi, m mV an f the -"fe to this very cnrioos paper. 


Chap. I. Rudiments, ig 

ally become developed into " thickset, long, and rather coarse 
" dart hairs," when abnormallj ncmriBhed near oid-etandJHg 
inflamed snrfaoes."" 

I am informed by Sir James Paget that often several members 
of a family have a few hairs in their eyebrows mueh longer than 
the others; so that eveii this slight peculiarity seems to be 
inherited. These hmrs, too, seem %a have their representatiyeB ; 
for in the chimpanzee, and in certain species of Macacna, there 
are scattered hairs of considerable length rising from the naked 
elrin above the eyea, and corresponding to our eyebrows ; similar 
long hairs project from the hairy covering of the superciliaiy 
ridges in Eome baboons. 

The fine wool-like hair, or so-called lanugo, with which the 
human fostns during the sizth month is thickly covered, offers a 
more curious casa It is first developed, during the fifth month, 
on the eyebrows and face, and especially round the mouth, 
where it is much longer than that on the head, A monstaehe 
of this kind was observed by Esohriohf ° on a female ftetus ; but 
this is not so snrprisii^ a circumstance as it may at first appear, 
for the two sexes generally resemble each other in all external 
characters dnrii^ an early period of growth. The direction and 
arrangement of the hairs on aJl parts of the fffital body are the 
same aa in the adnlt, bnt are subject to much variability. The 
whole surface, including even the forehead and ears, is thus 
thickly clothed; bnt it is asigaiScant fact that the palms of the 
hands and the soles of the feet are quite naked, like the inferior 
Burfaoes of all four extremities in most of the lower animals. As 
this can hardly be an accidental coincidence, the woolly cover- 
ing of the fcetus probably represents the first permanent coat of 
hair in those mammals which are bora hairy. Three or four 
cases have been recorded of persons bom with their whole bodies 
and faces thickly covered with fine long hairs ; and this strange 
condition is strongly inherited, and ia correlated with an abnor- 
mal condition of the teeth.'" Prof. Ales. Brandt informs ma that 
he has compared tte hair from the face of a man thus charac- 
'erised, aged thirty-five, with the lanugo of a fcetus, and finds it 
quite Bimilar in texture ; therefore, as he remarks, the case may 
he attributed to an arrest of dcTelopment in the hair, together 
with its continued growth. Many delicate ehildien, as I have 

•• Pagot, 'Lectures oa Sargical has teoently sent me an adilitional 

Pathology,' 1853, voL i. f. 71. casa of a falJier and bob, bora in 

*• Eschricht, ibid. b. 40, 47. ' Russia, with these peculiarities, i 

" Pse my ' Varration of Animals haie received drawinss of both frtm 

uid Plants abdet Domwiticsatiun,' Paris. 

^ iLp. 327. Prof. Alsx. BraDii 


20 Tile Descent of Man. Pabt I, 

lieen assured by a surgeon to a hospital for children, have tlieir 
backs covered by rather long silky hairs; aJid such cases pro- 
bably come under the same head. 

It appears as if the posterior molar or wisdoin-teeth were 
tendii^ to become rudimentary in the more oiviiised races of 
man. Those teeth are rather smaller than the other molars, aa 
is likewise the case with the corresponding teeth in the chim- 
panzea and orang ; and they have only two separate fangs. 
They do not out through the gums tUl about the seyenteenth 
year, and I haye been aBsured that they are much more liable fo 
decay, and are earlier lost than the other teeth ; but this is denied 
by some eminent dentists. They are also much more liable to 
vary, both in structure and in (he period of their development, 
than the other teeth." In the MeJanian races, on the other 
hand, the wisdom-teeth are usually furnished with Uireo 
wpai'ate fengs, and are generally sound ; they also differ irom 
the other molars in size, less than in the Caucasian races.*^ 
Prof. Schaaffhausen accounts for this difference between the 
races by " the posterior dental portion of the jaw being always 
" shortened" in those that are civilised," and this shortening may, 
1 presume, bo attributed to civilised men habitually feeding on 
soft, cooked food, and thus using their jaws less. I am informed 
by Mr. Brace that it ia becoming quite a common practice in the 
United States to remove some of the molar teeth of children, as 
the jaw does not grow large enough for the perfect development 
of the normal numbor." 

With respect to the alimentary canal, I have mot with an 
account of only a single rudiment, namely the vermiform append- 
age of the csecum. The caecum is a branch or diverticulum of 
the intestine, ending in a cul-de-sac, and is estremeJy long in 
many of the lower yegetable-feeding mammals. In the marsupial 
koala it is actually more than thrice as long as the whole body." 
It is sometimes prodaced into a long gradually-tapering point 
and is sometimes constricted in parts. It appears as if, in con- 
;e of changed diet or habits, the c»3um had become much 

« Dr. Webb, 'Teeth in Man ani 

from Florence, that he has lately 

tKe Anthropoid Apes,' as quoted by 

been studjing the last molar teeth 

Dr. C, Carter Biaie in 'Anthmpo- 

in tho different races of nwn, and 

legteal ReV«w,' Julj 1SS7, p. 29FP. 

*' Owen, 'Anatomy of Veite- 

that given in my teit, viz., that in 

bratM,' vol. iii. pp. 320, 321, anl 

the higher or eiviliaed races they 


are on Uie road towards atrophy or 

" 'OnthePrimitiTeFonnofthfl 

Skull,' Eng. translat. In 'Anthropo- 

" Owen, ' Anatomy of Verte- 

Icgieal Review," Oet. 186B, p. 4aa. 

brates,' voi, lii. pp. 41G, 434, 4il. 


1,'Bi? I. Rudiments. 2i 

Bhortened in Taiious animals, the vermiform appendase lieuig 
left as a mdiment of the shortened part. That this appendage 
ia a rudiment, we may infer from its small size, and from the 
evidence which Prof. Canestrini" haa collected of ite, variability 
in man. It is occasionally quite absent, or again is largely 
developed. The passage is sometimes completely closed for half 
or two-thirds of its length, with the terminal part consistii^ of 
a flattened soM expansion. In the orang tiiis appendage is long 
and convoluted: ia man it arises from the end of the short 
ciDcnm, and is commonly from four to five inches in length, 
being only about the third of an inch iu diameter. Not only is 
it nselesE, but it is sometimes the cause of death, of which fact 
I have lately heard two instances : this is due to small hard 
bodies, sucli as seeds, entering the passage, and cansii^ inflam- 

Ia some of the lower Qnadrumana, in the Lemuridffi and 
Carnivora, as well as in many marsupials, there is a passage near 
the lower end of the humerus, called the supra-condjioid fora^ 
men, through which the groat nerve of the fore limb and often 
the great artery pass. Now in the humerus of man, there is 
generally a trace of this passage, which is sometimes fairly well 
developed, being formed by a depending hook-hke process of 
bone, completed by a band of ligament. Dr. Struthers," who baa 
closely attended to the subject, has now shewn that this 
peculiaiity is sometimes inherited, as it has occnrred in a father, 
and in no less than four out of his seven children. When pre- 
sent, the great nerve invariably passes through it; and this 
clearly indicates that it is the homologue and rudiment of the 
supra-condyloid foramen of the lower animals. Prof Turner 
estimates, as he informs me, that it occurs in about one per cent 
of recent skeletons. But if the occasional development of this 
structure in man is, as seema probable, due to reversion, it is a 
retnm to a very ancient state of tiinga, because in the higher 
Qnadrumana it is absent. 

There is another foramen or perforation in the humeruSj 

" 'AnnuiiriQ della Soc. d. NaL' Feb. 15, 1873, and aootJier im- 

Modena, 1867, p. 94. ' portant paper, ibid., Jan. S*, 1863, 

'• M, C. Moi-tins (" De I'tlnittf p. 83. Dt. Knos, afl I am informed, 

Organiqne," in ' Rerne d«a Deax was the Krst anatomist who drc<r 

Moodes,' Juie 15, 1863, p. 16), and attention to this peculiar structure 

Hacfcel ('Generelle Motpliologie,' in m,in ; see hia 'Great Artists and 

B. ii, «. 278), have both remarked AnutomLsts,' p. S3. See also an im- 

on the singular fact of this rudi- ponaut memoir on this prooecs by 

- -- -ising deat"- "- "-■■' — ■" ''■" ' "■■i'"i" ■!■ 


22 The I^escent of Man. Past I 

present in man, whick may be called the inter- 
condyloid. This occurs, but not constantly, in various anthro- 
poid and other apes," and hkewiae in many of the lower animalR. 
It is remarkable that this x^tforation sooms to have been present 
in man much more frequently during ancient times than 
recently. Mr. Busk" has collected the following oyidence on 
this head: Prof. Broca " noticed the perforation in four and a 
" half per cent, of the aim-honea collected in the ' Oimetiere du 
" Sud,' at Paris ; and in the Grotto of Orrony, the contents of 
"which are referred to the Bronze period, as many as eight 
" humeri out of thirty-two were perforated ; hut this estraordi- 
" nary projwrtion, he thinks, might be due to the cavern haTing 
" been a sort of ' &mily vault.' Again, M. Dupont found thirty 
" per cent, of perforated bones in the caves of the Valley of the 
" Lesse, belonging to the Eeindeor period ; whilst M. Leguay, in 
" a sort of dolmen at Argenteuil, observed twenty-five per cent, 
"to be perforated; and M. Pmner-Bey found twenty-sii per 
" cent, in the same condition in bones from Vaureal. Nor should 
"it be left unnoticed that M. Pruner-Bey states that this con- 
" dition is common in Guaache skeletons." It is an interesting 
fact that ancient races, in this and several other cases, more 
frequently present structures which resemble those of the lower 
animals than do the modem. One chief cause seems to bo that 
the ancient races stand somewhat nearer in the long line of 
descent to their remote animal-lite progenitors. 

In man, the os coccyx, together with certain other vertebrfe 
hereafter to be described, though fucctionless as a tai!, plainly 
represent this part in other vertebrate animals. At an early 
embryonic period it is free, and projects beyond the lower 
estremities ; as may be seen m the drawing (Pig, 1.) of a human 
embryo. Even after birth it has been known, in certain rare 
and anomalous cases,'*'' to form a small external rudiment of a 
tail. The ob ooecyx is short, nsually including only four 
vertebrra, all ancbyloaed together : and these are in a rudi- 

'" Mr.SuOBoi^ MiTart,'Trana. " Qnatrefoges has lately collected 
vA. PhU. Soc' X887, p. 310. the svidenceon thi. snhjeot ' " 

•'" On the Caves of Gibraltar," ' " " " -"" " 
'Traasaot. Intemat. Congrass of 
PrehiBt. Arch.' Third SesBion, 1B69, 
p. 159. Prof. Wyman has lately 
thewD (Fourth Annual Report, Pea- casB, 
body Museuni,!871,p.20), that-' ' -'^- * 

Sm nneient moundB in the WesI 
ITultfd Statea, and la Florida, 
(rcquentlj oi-curi in the negro. 





In 1840 Fleis 

chmann ei- 

lited . 

a hui 

nan ftetu 

3 bearing a 

e tail. 

1 nlways the 




bodiea: and 


TitluxUy * 

...mined by 

! many ana 

WmistB pi 

■esent at tha 


of w. 


at Eriangen 

e Ma. 


in Nied 




logie, Deo 

ember! 87 1> 


Chat. 1. Rudiments. 23 

mentary eouditiMij fbr they consist, with the exception of the 
baaal one, of the centrum alone.'^ Thej are furnished with 
some small muscles; one of which, as I am informed by Prof- 
Tumer, has been espresal j described by Tlieile as a rudimentary 
repetition of the extensor of the tail, a muscle wMcli is bo 
largely developed ia many mammals. 

The spinal cord ia man extends only aa far downwards as t'na 
last dorsal or first lumbar vertebra; but a thread-like struc- 
ture (the jSJura ierminale) runs down the axis of the sacral part 
of the spinal canal, and even aloi^ the back of the coccygeal 
bones. The upper part of this filament, ns Prof. Turner 
informs, me, is undoubtedly homolt^ous with the spinal cord- 
but the lower part apparently consists merely of the pia matei; 
or vascular investing mombraue. Even in this case the os 
coccyx may be said to possess a vestige of so importout a 
structure as the spinal cord, though no longer enclosed within 
a bony canal. The following fa«t, for which I km also in- 
debted to Prof. Turner, shews how closely the os coccyx corre- 
sponds with the true tail in the lower animals : Luschka has 
recently discovered at the extremity of the coccygeal bones a 
VQtj peculiar convoluted body, which is continuous with the 
middle sacral artery ; and this discovery led Krause and Meyer 
to examine the tail of a monkey (Macacns), and of a cat, in both 
of which they found a similarly convoluted body, though not at 
the extremity. 

The reproductive system offers various rudimentary struc- 
tures; but these differ in one important respect frcan the 
foregoing cases. Here we are not concerned with the vestige of 
a part which does not belong to the species in an ef&cient state, 
but with a part efficient in the one sex, and represented in the 
other by a mere rudiment. Nevertheless, the occurrence of 
such rudiments is as difficult to explain, on the belief of the 
separate creation of ea«h species, as ia the foregoing cases 
Hereafter I shall have to recur to these rudiments, and shall 
shew that their presence generally depends merely on inheri- 
tance, that is, on parts acquired by one sex having been 
partially transmitted to the other. I will in this place only give 
some instances of such rudiments. It is well known that in the 
males of all mammals, inclading man, rudimentary mam mm 
exist. These in several instances have become well developed, 
and have yielded a copions supply of milk. Their essential 
identity in the two sexes is likewise shewn by their occasional 
sympathetic enlargement in both during an attack of the 

" Ow«D, ' On tbe Nature of Umhe,' 1849, p, 114, 


24 The Descent oj Man. Par™ i 

measles. The vesicala proslutica, which has been observed in 
many male mammals, is now imiversally acknowledged to he 
the homologne of the female uterus, together with the con- 
nected passage. It is impossible to read Leuckart's able 
deacription of this organ, and his reasoning, without admitting 
the justness of his conclusion. This is especially clear in th« 
case of those mammals in which the true female uterus 
bifurcates, for in the males of these the vesieula likewisa 
bjfurcatea." Some other rudimentary structures belonging ta 
the reproductive system might have been here adduced." 

The bearing of tie three great classes of facts now given is 
■nnmistakeable. But it would be superfluous fully to recapitulate 
the line of ajgament given in detail in my ' Origin of Species,* 
The homological construction of the whole frame in the members 
of the same class is intell^ible, if we admit their decent &om 
a common prngenitoi, together with their BTibsequent adaptation 
to iliversified conditions. On any other view, the similarity of 
pattern between the hand of a man or monkey, the foot 
horse, the flipper of a seal, the wing of a bat, &c., is uttijily 
inespUcable." It is no scientific explanation to assert tliat they 
have all been formed on the same ideal plan. With respect to 
development, we can clearly understand, on the principle of 

" Lenckart, in T.-dd's 'Cjclop. words) a mers metaphploal prin- 

of Anat." lI>49-52, vol iv. p. 1415. cipU, Lameljr, the preaerntion "in 

Ju maa this organ ia only from " its integrity of the mammalian 

three to SIX lines is length, but, " nature of ths animal." In only a 

like so many stber rudimeatniy few cases doea he discuss rndiments, 

part^ It js Tariahle in development and then only those parts which are 

as well as in other characters. partially rndimentaiy, soch as the 

" See, on this subject, Owen^ little hoots of the pig and ox, which 

pp. 675, 676, 706. ' shews dearly to ba of seriice to the 

" Prof. Sianconi, in a recently animal. It Is uafortuoate that he 

published work, illitstvated by ad- did not consider such eases as the 

mirable engravings ('La Th«>rie minute teeth, which never cut 

Sarwinianne et la criaticn dite in- through the jaw in the oi, or the 

dependante,' 1874), endeavoars to mamma of male quadrupeds, or the 

show that homological fitructurea, in win^ of certain beetles, eiiating 

the above and other cases, can be under the soldered wing-covers, or 

folly eiplaiued on mechauical priu- the vestiges of the pistil and stamens 

oiples, in accordance with their ases. in various flowers, sud many other 

Xo one has shewn so well, how ad- such cases. Althongh £ greatly 

mltably such structures are adapted admire Prof, Bianconi's work, jet 

for their final purpose; and this the belief now held by most natural- 

adaptatioa can, as I believe, be ists seems to me left unshaken. 

In considering the wing of a bat, he eiplioable on the principle of merf 
bliagsforward(p. 213)what3ppeju's adaDtation. 


C'liAP. T. Rudiments. 25 

variations Bupervening ai a rather late embijotiio period, and 
being inherited at a correspondiag period, how it is that the 
embryos of wonderfully different forms should still retain, morn 
or less perfectly, the structure of their common progenitor. 
No other explanation has ever been given of the marvellous fact 
that the embryos of a man, dog, seal, tat, reptile, iSc, can at first 
hardly bo distinguished from each other. In order to understand 
the existence of rudimentary organs, we have only to suppose 
that a former progenitor possessed the parts in question in a 
perfect state, and that under changed habits of life they became 
greatly reduced, either from daipla disnse, or through the natural 
selection of those iudiTiduals which were least encumbered with 
a superfluous part, aided by the other means previously in- 

Thus we can understand how it has come to pass that man and 
ail other vertebrate animals have been constructed on the same 
general model, why they pass throngh the same esrfy stages ot 
development, and why they retain certain rudiments in common. 
Consequently we ought frankly to admit their community of 
descent; to take any other view, is to admit that our own 
fitnictnre, and that of all the animals around us, is a mere snare 
laid to entrap our judgment. This conclusion is greatly 
utrengthened, if we look to the members of the whole animal 
series, and consider the evidence derived from their affinities 
or classification, their gec^aphical distribution and geolo- 
gical succession. It is only onr natural prejudice, and that 
arrogance which made onr forefathers declare that they were 
descended from demi-^ods, which leads us to demur to this 
conclusion. But the time will before long come, when it wiU be 
thought wondorftil that naturalista, who wero well acquainted 
with the comparative structure and development of man, and 
other mammals, should have believed that each was the worS 
of B eoparato act of creation. 


The Descent of Man 

Vsriability of bodj and mind in roan — ^nhsllt^nec — Canses of variability 
— LawB of vaiiatjou die same m niin as in the lower animals— Direct 
action of the conditiooB of life— Effects of the increased nse and disnu 
of parts — Ari-ested development — Reversion — Correlated vanation — 

dominant animal in the world — Importance of his corporeal etiueture— 

of structure— ^Decrease in size of the canine teeth — Increased size and 
altered Bha]« of the sknll— Nakedness— Absence of a tail— Defencelcia 
condition of man. 

It is manifest that man is now subject to much variability. 
No two indiYiduals of the same race aro quite alika We may 
compare miUioBa of fac«s, and each will bo distinct There ib 
an equally great amount of diversity in the proportions aad 
dimensions of the various parts of the body ; the lei^h of the 
legs being one of the most variable points.' Although in some 
quarters of the world an elongated skull, and in other quarters 
a short skull prevails, yet there is great diversity of shape even 
within the limits of the same race, as with lie aborigines of 
America and South AuBtra,Iia — the latter a race " probably as 
" pure and homogeneous in blood, cnetocis, and lai^age as any 
" in existence" — and even with the inhabitants of so confined 
un area as the Sandwich Islands.' An eminent dentist assures 
me that there is nearly as much diversity in the teeth as in the 
features. The chief arteries so frequently run. in abEOimal 
courses, that it has been found useful for surgical purposes to 
calculate from IMO corpses how often each, course prevails.' 
The muscles are eminently variable: thus those of the foot 
were found by Prof. Turner' not to be strictly alike in any two 
out of fifty bodies ; and in some the deviations were considerable 

' 'Investigations in Military and 

HQiiey, in Lyell's 'Antiquity 

Authropolog. Statistics of American 

Man,' 1863, p. 87. On the Sai 

Soldiers,' by B. A. Gould, 1869, p. 

wich Islanders, Prof. J. Wjmi 

'Observations on Crania," Best 

" With respect to tl.e "Cranial 

18S8, p. 18. 

forms of the Americau aboiiEines," 

' 'Anatomy of the Arteries,' 

see Dr. Aitten Meigs in ' Proc. 

K. Quain, Preface, vol. i. 1844. 

Acad. Nat. Scl.' Philadelphia, May, 

' "Transact. Royal Soc Ed 

tci^h,'Tol. «lv. pp. 17-., 1811. 


UBip. II. Manner of Development. 

lie adds, tlmt tte power of performing the appropriate movo- 
ments Enaat have been modified in accordance with the eeTei-al 
deviations. Mr, J. Wood has recorded' the occurrence of 295 
muBcular variations in thirty-six subjects, and in another set of 
the same number no less than 558 Tariations, those occurring on 
both sides of the body beir^ only reclioued as one. In the last 
Bet, not one body out of the thirty-Bii was " found totally 
" wanting in departures from the standard descriptions of the 
" muscular system given in anatpmical test books." A single 
body presettted the extraordinary number of twenty-five distinct 
abnormalities. The same ' muscle sometimes varies in many 
ways: thus Prof. Macalister describes' no less than twenty 
distinct variations in the yaimaris aeeessOTius. 

The famous old anatomist, "Wolff/ insists that the internal 
viscera are more variable than the esterual parts : NidUipartir 
cula est qux non aliler et aliter in aliis se habeat hommibus. He 
has even written a treatise on the choice of typical examples ot ■ 
the viscera for representation. A discussion on the beau-ideal 
of the liver, lungs, kidneys, &o., as of the human face divine, 
sounds strange in our ears. 

The variabihty or diversity of the mental faculties in men ol 
the same race, not to mention the greater differences between 
the men of distinct races, is so notorious that not a word need 
hore be said. So it is with the lower animals. All who have 
had charge of menageries admit this feet, and we see it plainly 
in our dogs and other domestic animals. Brehm especially 
insists that each individual monkey of those which he kept tame 
in Africa had its own peculiar disposition and temper; he men- 
tions one baboon remarkable for iU h^h jctelhgence ; and the 
keepers in the Zoological Gardens pointed ont to me a monkey, 
belonging to the New World division, equaUy remaikable for 
intelligence. Eengger, also, insists on the diversity in the 
various mental characters of the monkeys of the same species 
which he kept in Paraguay ; and this diversity, as he adds, is 
partly innate, and pan^ the lesnlt of the mauuei in which they 
have been treated or educated.* 

Ihaveelsewhere'sofullydiBcussedthesubject of Inheritance, i 
that I need here add hardly anythmg. A greater number of 

> 'Pi-oo. Royal Sue.' 1887, p. ■ Bl«hm, 'Tiiierleben,' B. i. s. 

1. 1868, p. Itl. 

■ 'Act. Ajad. St. PetarBbuig, 
1376, pait ii. p. 217 


25 The Descent of Man. FAarl 

facts hiiTH beun collected with respect to the tiansmission of the 
most trifling, as woll as of the moat important characters in 
man, than in aaj of the lower animals; though the facts are 
copious enougii with respect to the latter. So in regard to 
mental qualities, their transmiBaion is manifest in our dogs, 
horses, and other doiaesfie animals. Besides special tastes and 
habits, general iatelligonea, courage, bad and good temper, &c., 
are certainly transmitted. With man we stc sunilar facts in 
almost every family; and we now know, through the admirabla 
labours of Mr. Galton,"'that genius which implies a wonderfully 
complex combination of high faculties, tends to bo inherited; 
and, on the other hand, it is too certain that insanity and deteri- 
orated mental powers likewise run in families. 

With respect to the causes of variability, we are in all cases 
■very ignorant; but we can see that in man as in the lower 
animals, they stand in some relation to the conditions to which 
■ each species has been exposed, during several generations. 
Domesticated animals vary more than those in a state of nature; 
and this is apparently due to the diversified and changing natui* 
of the conditions to which they have been subjected. In this 
respect the different races of man resemble domesticated animals, 
And so do the individuals of the same race, when inhabiting a 
very wide area, like that of America, We see the influence of 
diversified conditions in the more civilised cations; for the 
members belonging to different grades of rant, and following 
different occupations, present a greater range of character than 
do the members of barbarous nations. But the uniformity of 
savages has often been exa^erated, and in some cases can hardly 
be said to exist." It is, nevertheless, an error to speak of man, 
oven if we look only to tho conditions to which he Las lieen 
exposed, as "far more domesticated"" than any other animal. 
Some savage races, such as the Anstralians, are not exposed to 
more diversified conditions than are many species which have 
a wide rangft In another and much more important respect, 
man differs widely from any strictly domestieated animal ; for 
his breedii^ has never long been controlled, either by methodical 
or unconscious selection. No race or body of men has been so 


'Heraiilfjy Genius: an In- 

•' man had sD oral viesge with (ina 

f into :ta Laws and CoDse- 

" features, and another was quit* 

icM,' 1869. 

"Mongolian in breadth and pro- 

Mr. Bates remarks (' The N;,tli- 

■' minence of cheek, spread of nos- 

t on the AmMofls,' ISeS, Toi. ii. 

"trils, and obliquity of eFes." 

i9). with respect to the Indiana 

le same Sonth American tribe. 

thro]>olog.' lioe. ttuitolal., iHi5, |i 

two of them wen at all .imilar 


Ihfl aiuipe of the liead; <-uf 


Chap. II. Manner of Development. 29 

completely subjugated by otlier men, as tliat certain individuals 
should be preserved, and thus imconseiously selected, from some- 
how escelling in utility to their masters. Kor have certain 
malo and female individuals been intentionally picked oat and 
matched, except in tie well-known case of the Prussian grena- 
diers; and in this ease man obeyed, as might have been ex- 
pected, the law of metliodical selection ; for it is asserted ttiat 
many tall men were reared in the Tillages inhabited by the 
grenadioTE and their tall wives. In Sparta, also, a form of aelec- 
tion was followed, for it was enacted that all children should be 
examined shortly after birth ; the woll-formed and vigorous 
being preserved, the otJiers left to perish." 

If we consider all the races of man as fonninga single species, 
bis range is enormous; but some separate races, as the Americans 
and Polynesians, have very wide ranges. It is a well-known 1 
law that widely-ranging species are much more variable than I 
species witb restricted ranges ; and the variability of man may 
vrith more trutti be compared witb that of widely-ranging species, 
than with tbat of domesticated animals. 

Not only does variabiUty appear to bo induced in man and 
the lower animals by the same general causes, but ia botti the 
same parts of the body are affected in a closely analc^ous 
manner. This, has been proved in such full detail by Godron and 

" Mitford's 'History of Gr 


vigoor of their chiyren. The Gre- 

vol. i. p. 282. It appears nlso 


cian poet, Theogais, who iivfd 550 

a passage in Xenophon's 'Ms 

B.C, clearly saw how importaiit 

hilia,' B. ii. 4 (to which my 

selection, if carefully BpplleiJ, would 

tiaa has been called by tbe 


be for the improrement of mankind. 

J. N. Hoare), that it was a 


He saw, likewise, that weaith often 


checks the proper action of seiual 

that mea onght to select their 

selection. He thus writes : 

with » view to the health 


By reasonable rales, and choose a bi'eed 
For profit and increase, at any price ; 
Of a sonnd stock, withoat defect or vice. 
But, in the daily matches that we make, 
The price is everything : for money's sake, 
Men marry ; women are in marriage giian 
The charl or ruffian, that in wealth has thriveu, 
May match his of&piing with the proudest race : 
Thus everything is mix d, noble and base ! 
If then in outward manoe:', form, and mini. 
You find as a degraded, motley kind. 
Wonder no mote, my tVieni 1 the cause is plain. 
Anil to lament the conseqaeDce is yajQ.'* 
CTha WcvWs of J. Hwfcham Ftere, vol. u. 1872, p. 3340 


30 The Descent of Man. Pi^T L 

Qnatrefages, that I need here only refer to their works.'' Mon- 
Rtrosities, which graduate into slight yariations, are likewise so 
Eimilar in man and the lower animals, tbat the eauie classification 
and the same terms can be used for both, as has been shewn by 
Isidore Geoffroy St.-Hilaire." In my work on the Tariation of 
domestic animals, I have attempted to arrange in a rude fafibion 
tie laws of Tariation under the following heads : — Thedirect and 
definite action of changed conditions, as exhibited by all or nearly 
ill the individuals of the same species, varying in the same manner 
[inder the same circumstances. The effects of the long-continued 
disuse of parts. The cohesion of homologous parts. The 
variability of multiple parts. Compensation of growth ; but of 
this law I have found no good instance in the case of man. The 
the mechanical pressure of one part on another ; as of 
the pelvis on the cranium of the infant in the womb. Arrests of 
development, leading to the diminution or suppression of parts, 
reappearance of long-lost characters through reversion. 
And lastly, correlated variation. All these so-called laws apply 
equally to man and the lower animals; and most of them oven 
It would be superfluous here to discuss all of them ;"" 
al are so important, that they must be treated at con- 

The direct and definite aciiim of changed conditions. — This is a 
most perplexing subject. It cannot be denied thatchauged con- 
ditions produce some, and occasionally a considerable effect, on 
organisms of all kinds ; and it seems at first probable that if 
sufficient time were allowed this would be the invariable result. 
Biit I have failed tc obtain clear evidence in favour of this con- 
clusion ; and valid reasons may be urged on the other side, at 
least as far as the innumerable structures are concerned, which 
are adapted for special ends. There can, however, be no doubt 
that changed conditions induce an almost indefinite amount of 
fluctuating variability, by which the whole organisation is rend- 
ered in some degree plastic. 

lu the United States, above 1,000,000 soldiers, who served in 
the late war, were measured, and the States in which they were 

" ajdron, 'De TEspfece,' 1859, 

» I have fully discoseed tiess 

trm.ii.liyre3. Quatrefeges, 'Dnite 

\A-ws -m ray 'Variation of Anlmnls 

in l'lLi|>e<» Humaine,' 1861. Also 

Leetnres oa Aothropologj, given in 

vol. it. ciiap. iiiL Slid iiiii. M. J. 

P, Durani has lately (1868) pub- 


lished a valuable easay 'De Pln- 

'• 'Hisl. Gin. !l Part, des Ano- 

flu«Dce des Milieu,' &c He lays 

malies de I'OrganisatioD," in three 

much stress, in the oaae of placts, on 

Mlan:«s, tola. i. 1833. 


Cbap. II. Manner of Development. 31 

bom and reared were recorded." From this astoiiisMng numbor 
of obaerrations it is proved that iocai influences of some kind 
act directly on etatnie ; and ma further learn that " the Stat« 
"where the ph^ical growth has in great measure token place, 
" and the State of birth, which indicates the ano^try, seem to 
" esert a marked influence on the stature." For instance, it 19 
established, " that residence in the Western States, during the 
" years of growth, tends to produce increase of stature." On the 
other hand, it is certain that with sailors, their life delays growth, 
^& shewn " by the great difference between the Btaturos of soldiers 
" and sailors at the ages of seventeen and eighteen years." Mr. B. 

A. Giould endeavouied to ascertain the nature of the influences 
which thus act on stature ; but he arriTed only at negative results, 
nasnely, that they did not relate to climate, the elevation of the 1 
land, soil, nor even "in any controlling degree" to the abundance I 
or the need of the comforts of life. This latter conclusion is 
directly opposed to that arrived at by Villerme, from the statistics 
of the height of the conscripts in different parts of France. When 
we compare the differences in stature between the Polynesian 
chie& aud the lower orders within the same islands, or between 
the inhabitants of the fertile volcanic and low barren coral islands 
of the same ocean," or again between the j^'uegians on the eastern 
and western shores of their country, where the means of snbsis- 
tence are very different, it is scarcely possible to avoid the con- 
clusion that better food and greater comfort do influence stature. 
Bat the preceding statements shew how difficult it is to arrive 
Bit any precise result. Dr. Beddoe has lately proved that, with 
the inhabitants of Britoin, residence in towns and certain occupa- 
tions have a deterioratii^ influence on height ; and he infers that 
the result is to a certain extent inherited, as is likewise the case 
in the United States. Dr. Beddoe further beheves that wherever 
a " race attains its rnfi.yiiniiTn of physical development, it rises 
" behest in energy and moral vigour." " 

Whether estemal conditions produce any other direct effect 1 
on man is not known. It might have been espeeted that dif- 
ferences of climatewould have had a marked influence, in as much 
aa the lungs and kidneys are brought into activity under a low 

" 'InTeetigatioils in Military and 289. There is also a remarkable 

Anthrop. Stiti»tics,' ic. 18fi9, by diflereace tn apjieorancB between 

B. A. Gonld, p. 93, 107. 128, 131, the closely-allied Hindoos Inhabiting 
134, tho Upper Ganges and Bengal ; sea 

It For the Polyneeiana, see Prioh- Elphinstone'i ' History of India,' vol. 

ard'9 ' Phyaical Hirf. of Mankind,' i. p. 324. 

vol. V. 184T, p. 145, 283. Also " ' Memoin, Anthropolog. Soo. 

PodroD, ' Da I'Eepics,' torn. ii. p. vol. iii. 1867-69, pf . 561, 56±i, 567. 


32 The Descent of Man. Paet 1 

tamperature, and the Iitbt and Etin under a Ligli one." It wofl 
foiRierly thonglit that the colonr of the skin and the character 
of the hair were determined by light or heat ; and although it 
can hardly be denied that Gome effect is thus produced, ahnost 
all observers now agree that the effect has been verj small, even 
after exposnie during manj ages. But this subject will be uioro 
properly discussed when we ijeat of the diSerent races of man- 
kind. With om domeBlJo animals there are grounds for 
believing that cold and damp directly affect the growth of the 
hair ; but I have not met with any evidence on this head in the 

Effects of the intreased Use and Disuse of Par's.— It is well 
known that use strengthens the nmscles in the individual, and 
complete disuse, or the destruction of the proper nerre, weakens 
them. When the eye is destroyed, the optic nerre olton becomes 
atrophied. When an artery is tied, the lateral channels increase 
not only in diameter, but in the thiekness and strength of their 
coats. When one Wdney ceases to act from disease, the other 
increases in size, and does double work. Bones increase not 
only in thickness, but in length, from carrying a greater weight." 
Different occupations, habitnaiJy followed, lead to changed 
proportions in various parts of the body. Thusit wasascertained 
by tho United States Commission " that the legs of the sailors 
employed in the late war were longer by 0217 of an inch, than 
those of the soldiers, though the sailors were on an average 
shorter men ; whilst their arms were shorter by 109 of an inch, 
and therefore, out of proportion, shorter in relation to their 
lesser height. This shortness of the arms is apparently due to 
their greater use, and is an unespeoted result ; but sailors 
chiefly use their arms in pulling, and not in supporting weights. 
With sailors, the girth of the neck and the depth of the inalep 
are greater, whilst the ciroumference of the chest, waist, and 
hips is less, than in soldiers. 

Whether the several foregoing modifications would become 
hereditary, if the same habits of life were followed during many 
generations, is not kno\Fn, but it is probable. Eengger'^ attri- 
butes the thin legs and thick arms of the Payaguas Indians to 

» Dr. Brakenridge, 'Theoiy of 

Dr. Jneger, "Ueber das Un, 

Diathesis," 'Mediral Times,' June 19 

Md Joly 17, 1869. 

" I Lave given authorities for 

ischen Zeitjchrift,' B. v. Heft i. 

" ' Invest; jationa," Sic Br t 

thtsB several etatemeaH fu mj 

Goold, 1869, p. 288. 

'Variation of Aniiuals under Do- 

mestication,' vol. ii. pp. 237-300. 


Chap, II. Marnier of Development. 33 

KncceEBive generations having passed nearly their whole lives in 
uanoes, with their lower extremities motionless. Other writers 
have come to a aimilticconolusioiiiiinnalogoiis cases. According 
to Cranz,"* who lived for a long time with the I^uimaux, " the 
" natives believe that ingenuity and dexterity ia seal-catehing 
" (their highest act and virtue) is hereditary ; there is really 
" something in it, for the son of a eelehrated seal-catcher will 
"difitii^niEh himself, though he lost his father in childhood." 
But in this ca^se it is mental aptitude, quite as much as bodily 
structure, which appears to be inherited. It is asserted that 
the hands of English labourers are at bu^h larger than those ol 
the gentry." From the correlation which exists, at least in 
some cases," between the development of the extremities and of 
the jaws, it is possible that in those classes which do not labour 
much with their hands and feet, the jaws would be reduced in 
Bize from this cause. That they are generally smaller in refined 
and civilised men than in hard-working men or savages, is certain. 
But with savages, as Mr. Herbert Spencer" has remarked, the 
greater use of the jaws in chewing coarse, uncooked food, would 
not in direct manner on the" masticatory muscles, and on the 
bones to which they are attached. In infants, long before birth, 
the skin on the soles of the feet is thicker than on any other part 
of the body;'^ and it can hardly be doubted that this is duo 
to the inherited effects of pressure during a long series ol 

It is familiar to every one that watcbmaiers and engravers 
are liable to be short-sighted, whilst men living much out o( 
doors, and especially savages, are generally long-sighted." Short- 
sight and long-sight certainly tend to be inherited." The 
inferiority of Europeans, in comparison with savages, in eye- 
sight and in the other senses, is no doubt the accumulated and 
transmitted effect of lessened use during many generations ; for 
Eengger" states that he has repeatedly observed Europeans, 

" 'History of Greenland,' Eng. ('Sanitarv Memoirs of the War ol 

tranalflt. 1767, vol. i. p. 230. the Rebellion,' 1869, p. 530). taa 

" ' Intann»rria£e.' By Alei. proTed this to be the case ; and h* 

Walker, 1838, p. 377. aceonnts for il by the ordinaiy 

" * The Variation of Animals range of vision in sailor! being " re- 

under Domestication, vol. i. p. 173. " stricteJ to the length of the vessel 

■' 'Principles of Biology,' tol. t, " and the height of tbo masts." 

p. 455. "'The ■Variation of Animals 

" Paget. ' Lectures on Sargical nnder Domesticnlion," vol. i. p. 8. 

Pathology,' vol. ii. 1853, p. 209. "'^ugethiera vua Paraguay,' 

"It ia a singular and nnei- a, 8, 10. I have tad good opportuni- 

peeted fact that sailors ar* inferior ties for observing the eitraordinary 
power of eyesight in the Fuegiaos. 
Sue also Lawreaco {'Leotnres ou 


34 The Descent of Mtui. Pjet!. 

who had been brought up and spent their whole lives with tha 
wild Indians, who noTorthelesa did not equal them in the sharp- 
noKS of their senses. The same naturalist observes that the 
cavities in the skull for the reception of the Keveral sense-oi^ans 
are larger in the American aborigines than in Europeans; and 
this probably indicates a corresponding difference in the dirnen- 
Biona of the organs themselves, Blumenbaoh has also remarked 
on the lai^e size of the nasal cavities in the skulls of the 
American aborigines, and connects this fact with their remarkablj 
acute power of smell. The Mongolians of the plains of Northern 
Asia, according to Pallas, have wonderfully perfect senses ; and 
Prichard belietes that the great breadth of their skulls across 
the zygomas follows from their highly-developed sense-oi^ans,™ 
The Quechna Indians inhabit the lofty plateaux of Peru ; and 
Alcide d'Orbigny states" that, from continually breathing a 
highly rarefied atmosphere, they have acquired chests and lungs 
of extraordinary dimensions. The cells, also, of tto lungs are 
larger and more nomerous than in Europeans. Those observa- 
tions have lieen doubted ; but Mr. D. Forbes carefully measured 
many Aymaras, an allied race, living at the height of between 
10,000 and 15,000 feet; and he informs me" that they 
differ conspicuously from the men of all other races seen by bim 
in the circumference and length of their bodies. In bis table of 
measurements, the stature of each man is taken at 1000, and the 
other measurements are reduced to this standard. It is here 
seen that the extended arms of the Aymaras are shorter than 
those of Europeans, and much shorter than those of Negroes. 
The legs are likewise shorter ; and they present this remarkable 
peculiarity, that in every Aymara measured, the femur is acfually 
Bhorter than the tibia. On an average, the length of the femur 
to that of the tibia is as 211 to 252; whilst in two Europeans, 
measured at the same time, the femora to the tibiie were as 244 
to 230; and in three Negroes as 258 to 241. The humerus is 
likewise shorter relatively to the foreaim. This shortening of 
that part of the limb which is nearest to the body, appears to be, 
as su^ested lo me by Mr. Forbes, a case of compensation in 

Physiology,' &c, 1822, p. 404) on bach, vol. i. 1851, p. 311; lor the 

this same snbjeot. M. eiraud-Tealon stnteniEnl by Pallas, vol. ir. 1844, 

has receatlj collected (' Kevue <]?s p. 40T. 

Conis Scientifiquea,' 1870, p. 895) "Quoted by Prichard, 'Ke- 
alorgBandTBlnable body of evidence aearcbes into the Phys. Hiat. of Man- 
proving that the canse of short- kind,' vol. v. p. 4Sa. 
light, "C«jt iV iTstaS oasidtt, de ^ Mr. Forbes' valnable papat i< 
uria." now published in the 'Journal of 
. of Man- the Ethnological Sc.. of London, 
Blumen- new leriea, Tol. il 1870, p. 1S3. 


OnAP. U. , Manner of Development. 35 

Telatjofl with the greatly increased length of the trank. The 
Aymaras present some other singular poiats of structure, foi 
instance, the very small projection of the heel. 

These men ate so thoroughly acclimatised to their cold and 
lofty abode, that when formerly carried down hy the Spaniards 
to the low eastern plaiaa, and when now tempted down by high 
wages to the gold-washinge, they suffer a frightful rate of mor- 
tality. Neverthele^ Mr. Forbes found a few pure families 
which had surrived during two generations : and he observed that 
they still inherited their characteristic peculiarities. But it was 
manifest, even without measurement, that theee peculiarities 
had all decreased ; and ou measurement, their bodies were found 
not to he so much elongated as those of the men on the high 
plateau; whilst their femora had become somewhat lengthened, 
as h&d their tibife, although in a less d^ree. The sotnal 
measurements may be seen by consulting Mi. Forbes's memoir. 
From these obserratious, there can, I think, be no doubt that 
residence during many generations at a great elevatiou tends, 
both directly and indirectly, to induce inherited modiflcations 
in the proportions of the body.'" 

Although man may not have been much modified during 
the latter stages of his esiatenoe through the increased or de- 
creased use of x>arts, the facts now given shew that his liability in 
this respect has not been lost ; and we positively know that the 
same law holds good with the lower animals. Consequently we 1 
may infer that when at a remote epoch the progenitors of mai 
were in a transitional state, aud were changing from quadrupedi 
into bipeds, natural selection would probably have been greatlj 
aided by the inherited effects of the increased or diminished usi 
of the different parts of the body. 

Arrests of Development. — There is a difference between arrested 
development ajid arrested growth, for parts in the former state 
contiuue to grow whilst still retaining their early condition. 
Various monstrosities come under this head; and some, as a 
clefl-palate, are known to be occasionally inherited. It will 
suffice for our purpose to refer to the arrested brain-development 
of microcephalous idiots, as described in Vogt's memoir." 
Their skulfa are smaller, and the convolutions of the brain 
are less complex than in normal men. The frontal sinus, or the 

" Dr. WadtBns ('LandwUth- regionfl, have liar frames modi Hl-I. 
Khaft. Wochenbbtt,' No. 10, 18ti9) "'Mrai^a sur lea Micron*, 

hu lately pobliihed an inUrestiog phales,' 1867, pp. 50, 135, Ilia, 171, 

gisli, whidi live in mouotaiuuts 


36 The Descent of Man. fABil. 

pnijection over the eye-brows, is lai^ely devcloiied, and the jaws 
ore pn^cathois to an "effrayani" degree; su that tlese idiots 
somewhat i^semhle the lower types of mankind. Their in- 
telligence, and most of their mental faculties, are estremely 
feeble. Thoj camiot acquire the power of speech, and are 
wholly incapable of prolonged attention, but are much given to 
imitation. They are strong and remarkably active, continually 
gamboling and jumping about, and making gritaaces. They 
often ascend stairs on all-foura; ajid ai'e curiously fond of 
climling up furniture or trees. We are thns reminded of th« 
delight shewn by almost all boys in climbing trees; and thia 
f^ain reminds us how lambs and kids, originally alpine aciinals, 
delight to frisk on any hillock, hcwever smah. Idiots also 
resemble the lower animals in some other respects; thus several 
cases are recorded of their carefully smelling every mouthful of 
Ibod before eating it. One idiot is described as often using his 
mouth in aid of his hands, whilst hunting for lice. They are 
often filthy in their habits, and have no sense of decrau^; and 
several cases have been pnhhshed of their bodies being i»- 
markably hairy." 

fleuersion. — Many of the cases to be here given, might have 
been introduced imder the last heading. When a structure 
is arrested in its development, but still continues growing, 
•intil it closely resembles a corresponding strocture in some 
lower and adult member of the same group, it may in one sense 
be considered as a case of reversion. The lower members in a 
group give us some idea how the common progenitor was 
probably constructed; and it is hardly credible that a complex 
part, arrested at an early phase of embryonic development, should 
go on growing so as ultimately to perform its proper function, 
unless it had acquired such power during some earlier state of 
oiistence, when the present exceptional or arrested structure 
was normaL The simple brain of a microcephalous idiot, in as 
far OS it resembles that of an ape, may in this sense be said to 
offer a case of I'eversion.'' Thei'o are other cases which come 

" Prof. Lajeoct suma up tho pp. 48-51. Pinel has hIeo 

character of brute-like idiuts by striking case of hairiness 

calling them tlieroid ; ' Journal of idiot. 

Ueutftl Soiecct,' Jnly 1863. Dr. " In my ' Variation of 

Scott ('The Deaf aad Dumb," 2nd under Domestication ' (toI. i 

edit., 1870, p. 10) hoa often ob- I attributed the not rery ri 

served the imbecile Bmelliug their ofEUpemuiaerarymainniK i. 

(hod. See. on thb same subject, to reversion. I was led to 

Ksndiiey, ' Body and Mind,' 1870, 


Manner of Development. 

mo e et y tmd our p esen h ad of reverBion, Cottain 
i n. ur gna yoc imng ntli wer members of tlie group 

refore^ as I supposed, 

P of. Gcgenbaar (' Jenmschen 
hrift," B. V. Heft 3, e. 341X 
^^ di p tes Owen's ooQcluaioo. On the 
hnnd, aceoi'diDg to the opiaios 
sMaaixi by Di'. Giinther, on 
paddle of Ceratodua, which i» 
huciag given bo much milk that the provided vith articnlated bony rayt 
child was thus nourished. The pro- on both sides of a central chain of 
liability that the additional mammai bones, there seemsno great difficulty 
are due to reversion is -thas nuch in admitting that six or more digits 
weakened; nevertheless, it still on one side, or on both sides, might 
tr^ms to me probable, because two reappear throagh reTersion. 1 am 
pairs are often found symmetrically informed bj Dr. Zouteyeeii that 
on the breast ; and of this I myself there is a case on record of B man 
have received information in several having twenty-four fingers and 
cases. It is well known that some twenty-four toes 1 I was chiefly led 
Lemurs normally have two pairs of to the conclusion that the presence 
mammie on the breast. Five cases of snpernoroevary digits Qiight be due 
have been recorded of the presence to reversion from the fact that such 
of more thau a pair of mammie (of digits, not only are sti'ongly in- 
coarse rudimentary) in the male herited, but, as I then believed, had 
sei of manltind ; see 'Joarnal of the power of regrowth after ampo- 
Anat, and Physiology,' 1872. p. 5^, tation, like the normal digits of the 
for a ease given by Dr. Handyside, lower vertebrata. But i have ei- 
in which two brothers eihihited plained in the Second Edition of my 
this pecnliEirity ; see also a paper by Variation under Domestication why 
Dr. Bnrtels, in ' Reichert's and du I now place little reliance on the 
Bois Rcymond's Archiv.,' 1873, p. recorded cases of such regrowth. 
304. In one of the cases alluded to nevertheless it deserves notice, in 
by Pr. Bartels, a man bore five as much as arrested development 
mammie, one being medial and and reversion are intimately related 
placed above the navel; Meckel processes; that various stroctnres 
von Hemsbach thinks that this in an embryonic or arrested con- 
latter case i< illustrated by a dition, such as a deft palate, bifid 
medial mamma occnrrinj in ceriain uterus, &c., are frequently noconi- 
CheiroDtera. On the whole, wc ma? panied bj polydaotyllsm. This has 
been strongly insisted on by Meckel 
and Isidore GeoliroySt.-Hil«ire. Bnt 

early progenitors been provided with give up altogether the idea that 
more than a single pair. there is any reiatioD between the 

In the above work (vol. ii. p. IT), ' ' 
I also Bttribnted, though with much 
besit-ition, the freoufut cities of 

i byGo Qt^le 

38 Tlie Descent 0/ Man. Past I, 

to which man belongs, occasionally make their appearance in 
bim, though not fonnd in the normal humaa embryo; or, \*. 
normally present in the human embryo, they become abnornially 
developed, althougli in a manner which is normal in the lower 
members of the group. These remajrfcs will be rendered clearer 
by the following illustrations. 

In variouB mammals the uterus graduates from a double 
organ with two distinct orifices and two passages, as in the 
marEupials, into a single organ, which, is in no way double 
except from having a shght internal fold, as in the higher apes 
and man. The rodents exhibit a perfect series of gradations 
between these two extreme states. In all mammals the uterus 
is developed from two simple primitive tubes, the inferior 
portions of which form the comua ; and it is lE the words of 
Dr, Farro, " by the coalescence of the two comaa at their lower 
" extremities that the body of the uterus is formed in man ; 
" while in those animals in which no middle portion or body 
'' exists, the comua remain ununited As the development of 
" the uterus proceeds, the two comua become gradually shorter, 
" until at length they are lost or, as it were, absorbed into the 
" body of the ctems." The angles of the utenis are still 
produced into cornua, even m animals as high up in the scale as 
the lower apes and lemurs. 

Now in women, anomalous cases are not very infrequent, in 
which the mature uterus is furnished with comua, or is partially 
divided into two oi^ns; and such cases, according to Owen, 
repeat "the grade of concentrativo development," attained by 
certain rodents. Here perhaps we have an mstaiico of a simple 
arrest of embryonic development, with subsequent growth and 
perfect functional development ; for either aide of the partially 
double uterus is capable of performing the proper office of 
gestation. In other and rarer cases, two distinct uterine cavities 
aj'e formed, each havii^ its proper orifice and passage.^' No such 
stage is passed through during the ordinary development of the 
embryo; and it is difBcult to beheve, thongh perhaps not im- 
possible, that the two simple, minute, primitive tubes should 
iBow how (if such an expression may be used) to grow into two 

"See Dr. A. Furre's well-known brates,'™]. iii., 1868, p. 687, Pro 

article in the ' Cydopaidia of Ad.i- fessor Tumsr in ' Edinburgh Meii- 

toioy and Fhjsioiogy,' vol. v. 1659, ra! Joninai,' Ffibruary J865, 
. 643. Owen, ' iuatoiny of Vorte- 


Chap, II. Manner of Development. 39 

distinct uteri, each wifli a well-constrticted orifice and passage, 
and each furmshed witli numerous muscles, nerves, glaoda and 
vessels, if they had, not formerly passed through a similar course 
of development, as in the case of existing maistipials. No ono 
wiU pretand that so perfect a structure as the abnormal douhlo 
nteruB iu woman could be the result of mere chance. But the 
principle of reversion, by 'which a long-lost structure is called 
back iato existence, might eerve as the guide for its full dovelop- 
ttent, even after the lapse of an enormous interval of time. 
Professor Cauestrini, after discussing the foregoing and various 
38, arrives at the same conclusion as that just 
He adduces another instance, in the case of the malar 
bone,*" which, in some of the Quadrumana and other mammals, 
normally consisis of two portions. This is its condition in tte 
human fcetus when two months old ; and through arrested develop- 
ment, it sometimes remains thus in man when adult, more 
especially in the lower prognathous races. Hence Canestrini 
concludes that some ancient progenitor of man must have had 
this bone normally divided into two portions, which afterwards 
became fused together. In man the frontal bone consists of a 
single piece, hut in the embryo, and in children, and in almost 
all the lower mammals, it consists of two pieces separated by a 
distinct suture. This suture occasionally persists more or less 
distinctly in man after maturity ; and more frequently in ancient 
than in recent crania, especially, as Canestrini has observed, id 
those eihtuned from the Drift, and belonging to the braehyce- 
phalio typo. Here again he comes to the same conclusion as in 
the analogous case of the malar hones. In this, and other instances 
presently to be given, the cause of andent races approaching the 
lower animals in certain characters more frequently than do the 
modern races, appears to be, that tlie latter stand at a somewhat 

nlisti in Modeus,' lb6T ] B3 ndult eknlls he l9D temarke tfant 
Prof. Caoestrini gives fjtracts on il more frequeutlv otcurs in pio- 

; sutject Tra 

the 'two malar bones in several ma.are Tonno ld72 Al«o E. 

huniaii sntijeots and in certaiQ apes MoneUI Sopra nna raia uiomalia 

he cannot consider this di<positicn c1»ll osso milare Modena, IS?" 

ef the parts as simply accidentjl St 11 more lecentlv Gcuber hns 

Another paper on this same anoma T written a lamphlet on the division 

has been published bj Di SsMott f this bone 1 give these i eferencet 

In the 'Gaiictta delle CI n che beransB a rtviewer without say 

Turin, 1871, whew h says thit gronnds or «rrnplef has thrown 

traces rf the division niaj be de doubts or my itatements 


40 The Descent of Man. PjetL 

(creator distance in tho long line of descent from their early senii- 
(luman progenitors. 

Various otlier anomalies in mim, mare or less anali^oiis to the 
forefcomK, have been advanced by different autliors, as cases of 
fCTersion; bnt tiieso seem not a little doubtful, for we httTO tfl 
descend estremely low in the mammalian fieries, before we find 
such Btrnetures normally present." 

In man, the canine teeth are perfectly efficient instrnments fo; 
mastication. But their true canine character, as Owen" re- 
marks, " is indicated by the conical form of the crown, which 
" terminates in an obtnse point, is convex outward and flat or 
sub-concave within, at the base of which surface there is a 
feeble prominence. The conical form is 
Melanian races, especially the AustraliaiL The e 
deeply implanted, and by a stronger fang than the ineisors." 
Nevertheless, this tooth no longer serves man as a special weapon 
for tearing his enemies or prey ; it may, therefore, as far as its 
proper function is concerned, be considered as mdimentary. In 
every large collection of human skulls some may be found, as 
Hackel" observes, with the canine teeth projecting considerably 
beyond the others in the same manner as in the anthropomorphous 
apes, but in a less degree. In these cases, open spaces between 
the teeth, in the one jaw are left for the reception of the eaninoa 
of the opposite jaw. An interspace of this kind in a Kaffir 
skull, figured by Wagner, is Eurprisingly wide." Considering 
how few are the ancient skulls which have been examined, 
compared to recent skulls, it is an interesting fact fiat in at 
least three cases the canines prejeet largely; and in tho Nanletf o 
jaw they are spoken of as enormous."' 

bj Isid. GsbflioT St..Hilaire, 'Hist, stance, in shnrteniag snd aii!iplifying 

dea AnomnUes,' torn. iii. p. 437. the course of development? AdJ 

A i-eviewer (' Journal of Anat. and again, why should not injurions ab- 

Physiology,' 1871, p. 366) blames normalities, such as atrophied op bj-- 

me much for not having discussed pertjophied parts, which have no 

says that, according to my theory, *i ' Anatomy of Vertebrates,' 

" CTcry transient condition of an iii. 1868, p. 323. 

" organ, during its development, is *' 'Generelle Morphologie," If 

" not only a means to an end, but B. ii. s. civ. 

" once was an end in itself," This " Cai'l Vogt's 'Lectures on M 

does not seem to me necessarily to Eng. tr^nslat. 1864, p. 151. 

hold good. Wby should not varia- J* C. Carter Blake, on i 

tious occur during an early period from La Naulette, 'Anthropo 

of development, having no relaUon Review," 1867, p. 296. Scha 

•a reversion; yet snoh v.-iriations hausen, ihid- 1K68, p. 426. 

might be preserved and aecurauUted, 


Chap, II. Maimer of Development. 41 

Of tbo antliropcunorphous apes the males alone have their 
canines fully doYeloped ; hut in, the female gorilla, and in a less 
d^jeo in tho female orang, these teeth project considerably 
beyond the others; therefore the fact, of which I have been 
aaaured, that ■women sometimes ha^e conaiderahly projecting 
canines, is no serious objection to the belief that their occasional 
great development in man is a case of reTeraion to an ape-lite 
prc^enitor. He who rejects with scorn the belief that the shape 
of his own canines, and their occosional great development in 
other men, are due to our early forefathers hayiog been pro- 
vided with these formidable weapons, will probably reveal, by 
Hneering, the line of Ms descent. For though he no longer 
intends, nor has the power, to use these teeth as weapons, he will 
unconsciously retract hia " snarling muscles" (thas named by 
Sir C. Bell)," so as to ospoae them ready for action, Uke a d<^ 
prepared to fight 

Many muscles are ooeasionally developed in man, which are 
proper to the Quadmmana or other mammals. Professor 
Vlacovich" examined forty male subjects, and found a muscle, 

called by him the 
others there was a 
a the remainii^ 

isohio-pubic, in nineteen of them ; in three 
ligament which represented this muscle; and 
eighteen no trace of it. In only two out o( 
thirty female subjects was this muscle developed, on both sides, 
but in three others the rudimentary ligament was present. This 
muscle, therefore, appears to be much more common in the 
male than in the female ses; and on the lielief in the descent 
of man from some lower form, the fact is intelligible ; for it 
has been detected in several of the lower animals, and in all 
of these it serves exclusively to aid the male in the act o£ 

Mr. J. Wood, in his valuable series of papers," has minutely 
described a vast number of muscular variations in man, which 
resemble normal structures in the lower animals. The muscles 

" ' The Anatomy of Eipressiou,' pp. 241, 242 ; vol. iv. 1867, p. 544 ; 

1844. pp. 110, 131. yo), ivi. 1868, p. 524, 1 may here 

" Quoted by I'rof. Caneatrini in sdJ that Dr. Murie and Mr. St. 

the 'Annuario ' &c, 1367 p 90 Geoi'ge Mivart haie shewn in their 

'^ These p^peli deserve careful Memoir on the Lemuroidea (* Trac- 

ituif by any one who desires to sact, Zoolog. Sot' vol. viL 1869, 

learn how frequently our mn-ck. p '>i\ how Bitraordinarily variable 

vary, and m varying come to re- some of the muscle* are In these 

cemble those of the Quadrnmana animals, the lowest memben of the 

Tlie foIlowinE refereacea lelate to Primates. Gradations, also, in the 

the few points t«uched on m my mnscles leading to structars fonnd 

t*it ' Proo Royal Soc vol iiv in animals still lower in the scnie, 

1865, pp 37»-3B4, vol I» 1868, *.-a nnmenns in the Lemncoidea. 


42 The Descent of Man. Taet 1. 

which closely reserable those regularlj present in our nearest 
(lilies, the Qufldrnmana, aro too numerous to be here even 
BpcciSed. In a singla male subject, having a strong bodily 
frame, and woU-formod slinll, no less than seren musculw Taria- 
tions were observed, all of whicli plainly represented muscles 
proper to various Idiids of apes. This man, for instance, had on ■ 
both sides of his neck a true and powerful " levator davieulm," . 
Buch as JR found in all kinds of apes, and which is said to occur 
in about one oat of sixty human subjectB." Again, this man 
had " a special abductor of the metatarsal bone of the fifth 
" digit, such as Professor Huxley and Mr, Flower have shewn 
" to exist uniformly in the higher and lower apes." I will give 
only two additional eases; the acromio^silar muscle is found 
in all mammals below man, and seems to be correlated with a 
inadmpedal gait," and it occnrs in about one out of sixty 
human subjects. In the lower extremities Mr. Bradley " found 
an abduclor ostis metatarsi quinti in both feet of man ; this muscle 
had not up to that time been recorded in mankind, bet is 
always present in the anthropomorphous apes. The muscles of 
the hands and arms — parts which are soeminentlycbaraoteristic 
of man— are extremely liable to vary, so as fo resemble the 
corresponding muscles in tbe lower animals.™ Such resem- 
blances are either perfect or imperfect; yet in the latter case 
they aro manifestly of a transitional nature. Certain variations 
are more common in man, and others In woman, without our 
being able to assign any reason. Mr. Wood, after describing 
numerous variations, makes the following pregnant remark. 
" Notable departure from the ordinary type of the muscular 
" structures mn in grooves or directions, which must be taken 
" to indicate some unknown iactor, of much importance to a 
" comprehensive knowledge of general and scientific anatomy.'™ 

" Sep also Prof. MacalLster in 

able ca» of variation in the Umu> 

' Proc R. Irish Academy,' vol. i. 

fiexor poinds /on^uj, adds, "Thi» 


=8, p. 134. 

" remarkable eiample shews thai 

" Mr. Champnep id 'Joornal of 
nt. aad Thys.' hW, 1B71, p. 178. 
>' 'Journal of Aoat. and Fhyj.' 

" arcangement of (endons 0/ thumb 

" and fingers characteristic of the 


J, 1872, p. 421. 

" case Ehonld be legarded «s a 


i taholated his observations, and 

" macaque passing upwards into a 


i> that muMnlar abnortdalitiea 

" TDsn, or a man passing downwards 

" into « macaque, or as a congenital 

Dndiy, in the face, thirdly, in the 

" freak of nature, I cannot uider. 


t, &c 

" take to say." It is satisfactory 

" The Rev. Dr. Haughton, after 

to hear so capabls an anatomist, 


■iBg ('Proo. K. Irish Academy,' 

and so embilttred an opponent ol 

.e 37. 186+, p. 715) a remark- 

PTolutionism, admitting even th( 


CuAr. 11. Manner of Development. 43. 

That tliis unknown factor is reversion to a fonner state oi 
existence maj be admitted as in the highest degree probable " 
It is quite incredible that a man should through mere accident 
abnormally resemble certain apes in no less than seven of his 
muscles, if there had been no genetic connection between them. 
On the other hand, if man is descended from some ape-like 
creature, no valid reason can be asBigned why certain muscles 
should not suddenly reappear after an interval of many thou- 
sand generations, in the same manner as with horses, asses, and 
mules, dark-coloured stripeB suddenly reappear on the legs, 
and sboulderB, after an interval of hundreds, or roore probably 
of thousands of generations. 

These various cases of reversion are so closely related to those 
of rudimentary Organs given in the first chapter, that many of 
them might have been indifferently introduced either there or 
here. Thus a human uterus furnished with cornua may be said 
to represent, in a rudimentary condition, the same organ in its 
normal state in certain mammals. Some parts which are rudi- 
mentary in man, as the os coccyx in both sexes, and the mammto 
in the male sex, are always present; whilst others, such as the 
Bupracondjloid foramen, only occasionally appear, and therefore 
might have been introduced under the head of reversion. Tliese 
several reversionary structures, as well as the strictly rudi- 
mentary ones, reveal the deseent of man from some lower form 
in an unmistakable manner. 

CarrAaUd Variation.— Jn man, as in the lower animals, many 
structures are so intimately related, that whai one part varies 
so does another, without our being able, in most cases, to assign 
any reason. We cannot .say whether the one part governs the 
other, or whether both are governed by some earlier developed 

possibility of either of his first pro- 

closely the vati^Uons resemble the 

positions. Pmf. Macalister W also 

normal muscles of the lower «ni- 

dMcril>ad ('P™. R. Irish Acad.' 

mais. He sum. up by remarking, 

yoL 1. 1881, p. 138) vai^iationa in 

" it will be enough for my purpose 

" if 1 have succeeded in shewing 

from their relatioDB to the same 

mosde iD the Quadrumaua. 

" when octurring as vai-ieties in the 

" Sinus the first edition of this 

" human subject, tend (o. eihibit ii- 

1»ok npu«aied, Mr. Wood h»s pub- 

" a sufficienlly marked manner what 

lished another memoir io the 'PhiL 

" may be considered as proofs and 

Tranaactions,' 1870, p. 83, on the 

" oamples of the tiarivinian prii*. 

varieties of the muscles of the haman 

" ciple of reversion, or law of iu- 

neck, shoulder, and chest. He here 

shews bow eitremely variable these 

muBolei art. nnii how often and how 


The Descent of Man. PaaT I 

part Various monstrosities, as I. Geoffroj repeatedly insists, aw 
thus intimately cocnectad. Homologous stnicturea are par- 
ticnJarly liable tw change ti^ethor, as ive see on the opposite 
sides of the body, and in the upper and loirei extremities, 
Meckel long ago remarked, that when the muscles of the arm 
depart from their proper type, they almost always imitate those 
of the leg; ajidso, conTersely, witJi the muscles of tie lege. The 
oi^na of sight and hearing, the teeth and hair, the colour of the 
skin and of the hair, colour and constitution, are more or less cor- 
related." Professor Sehaaffhausen flret drew attention to the 
relation apparently existing between a muscuiar frame and 
the strongly-pronounced supra-orbital ridges, which are so 
characteristic of the lower races of man. 

Besides the variations which can be grouped with more or 
less probability under the foregoing heads, there is a large class 
of variations which may be provisionally called spontaneous, for 
to our ignorance they appear to arise without any exciting 
cause. It can, however, be shewn that such variations, whether 
consisting of slight individual differences, or of Etrongiy-marked 
and abrupt deviations of structure, depend much more on the 
constitution of the organism than on the nature of the condi- 
tions to which it lias been subjected.'' 

Haie 0/ Increase. — Civilised populations have been known 
under favourable conditions, as ia the United States, to double 
their numbers in twenty-five years ; and, according to a calcula- 
tion byEuler, this might occur in a little over twelve years." At 
the former rate, the present population of the United States 
(thirty millions), would in 657 years cover the whole terrac[ueoua 
globe so thickly, that four men would have to stand on each 
square yard of surface. The primaiy or fundamental check to 
the continued increase of man is the difficulty of gaining 
subsistence, and of living in comfort. We may infer that this is 
the case from what we see, for instance, in the United States, 
where subsistence ia easy, and there is plenty of room. If such 
means were suddenly doubled in Great Britain, our number would 
bo quickly doubled. With civilised nations this primary check 
acts chiefly by restraining marriages. The greater death-rate of 
infants in the poorest classes ia also very important ; as well as 

" ThB 

Buthoritiea for tliese seve- 

■ my ' Variation of Animals 

and PI. 

ral jUtei 

. of Animsla under Do- 

" See tlie ever memom 


a' vol. iL pp. 320-a35. 

on the Principle of Popn 

" Thi» 

whole solyect hsa beei 

1 the R«v. T. Ualthnf., vol. 



in olwp, irili. ^l. ii. 0; 

r e, 517. 


OttiP. II. Manner of Developitient. 45 

the greater mortality, from various diseases, of tte mba'cdtanle o( 
crowded and miserable honsea, at all agos. The effects of severe 
epidemics and wars are Boon counterbalanced, and more thaji 
coontorbalanced, in nations placed nnder fiiTourable conditions. 
Emigration also comes in aid as a temporarj check, bat, with 
the extremely poor classes, not to any great extent. 

There is reason to suspect, as Malthus has remaiked, that the 
rcprodnctive power ia actually less in barbaious, than in civilised 
races. We know nothing positively on this head, for with 
savages no census has been taken; but from the oonenrtent 
testimony of missionaries, and of others who have long resided 
with such people, it appears that their families are usually small, 
and large ones rare. This may ba partly accounted for, as it is 
believed, by the women Budding their infents during a long 
time ; but it is highly probable that savages, who often suffer 
much hardship, and who do not obtain so much nutritious food 
as civilised men, would be actually less prolific I have shewn 
in a former work,'*that all om; domesticated quadrupeds and 
birds, and aU our cultivated plants, are more fertile than the 
correepondii^ species in a state of nature. It is no vahd 
objection to this conclusion that animals suddenly supplied with 
an excess of food, or when grown very fat ; and that most plants 
on sudden removal from very poor to very rich soil, are 
rendered more or less storila We might, therefore, espect that 
civilised men, who in one sense are highly domesticated, would 
be more prolific than wild men. It is also probable that the 
increased fertihty of civilised nations would become, as with onr 
domestic animals, an inherited character : it is at least known 
that with mankind a tendency to produce twins runs in 

Notwithstanding that savages appear to bo less proliSc than 
civilised people, they would no doubt rapidly increase if their 
numbers were not by some means rigidly kept down. The San- 
tali, or hill- tribes of India, have recently afforded a good illustra- 
tion of this fact ; for, as shewn by Mr. Hunter," they have 
increased at an extraordinary rate since vaccination has been 
introduced, other pestilences mitigated, and war sternly repressed. 
This increase, however, would not have been poBsibie had not 
■ these rude people spread into the adjoining districts, and worked 
for hire. Savages almost always marry ; yet there ia some 
prudential restraint, for they do not commonly marry at the 

" 'Variation of 

Auimala anil Fureign Meiliia^Chirurg. Berieiv, 

Piaafa under Domeal 

:ication,' vol. iL Jnly, ia33, p. 170. 

pp. 1I1-]13, 163. 

" ' The Annals of Rural Benjal, 

« Mr. Sertgivlti, 

'Bntisli and bj- W. W. HnnUr, ISUt p. 250. 


46 The Descent of Man. I'al.t L 

earliest possibie age. The joung men are often required to shew 
that they cim support a wife ; and they generally have fiibt to 
earn the price with which to purchaso her from her patents. 
With savages the difficulty of obtaining BTibsiatance occasionally 
limits their number in a much more direct maauor thaa with 
civilised people, for all tribes periodically suffer from severe 
famines. At such, times savages are forced to deTour much had 
food, and their health can hardly fall to be injured. Many 
accounts have been published of their protruding stomachs and 
emaciated limbs after and during famines. They are then, also, 
compelled to wander much, and, as I was assured in Australia, 
their infants perish in large numbers. As famioes are peiiod- 
ical, dopendiiig chiefly on estreme seasons, ail tribes must 
fluctuate in number. They cannot steadily and regularly 
increase, as there is no artificiai increase in the supply of food. 
Savages, when hard pressed, encroach on each other's territories, 
and war is the result; but they are indeed almost always at war 
with their neighbours. They are hablo to' many accidents on 
land and water in their search for food; and in some countries 
they suffer much from the laiger beasts of prey. Even in 
India, districts have been depopulated by the ravages o_' 

Malthus has discussed these several checks, but he does not 
lay stress enough on what is probably the most important of all, 
namely infajiticide, especially of female infants, and the habit of 
])rocuring abortion. These practices now prevail in many 
quarters of the world; and infenticido seems formerly to have 
prevailed, as Mr. M'Lennan" has shewn, on a still more extensive 
scale. These practices appear to have originated in savages re- 
cognising the difSculty, or rather the impossibility of supporting 
all the infants that are born. Licentiousness may also be added 
to the foregoii^ checks; but this does not follow from failing 
means of subsistence; though there is reason to believe that in 
some eases (as in Japan) it has been intentionally eneourag^^d 
as a means of keeping down the population. 

If we look tiack to an extremely remote epoch, before man had 
arrived at tie dignity of manhood, he would have been guided 
more by instinct and less by reason than are the lowest savages 
St the present time. Our early semi-human progenitors would 
not have practised infanticide or polyandry; for the instincts ol 
the lower animnla are never so perverted " as to lead them ro- 

' Primitive Marriaga.' 1865. ments as followe 

A wrilcrin tha ■Spectator' " Mr. Darwin fin, 


CiiiP. it. .Mantief of Develop.ment. 47 

.gularij to destroy their own offspring, 01 to be quitD devoid of 
jealousy. There would have been no prudential restraint ftum 
marriage, and the aeies would have freely united at an early age. 
Heuce the progenitors of man would have tended to increaao 
rapidly; but checks of some kind, either periodical or constant, 
must bave kept down their numbers, even more eererely than 
with existing savages. What the precise nature of theso checks 
were, we cannot say, any more than with most other animals. 
We know that horses ami cattle, which are not extremely prolific 
animals, when first turned loose in South America, increased at 
an enormous rate. The elephant, the slowest breeder of all 
known animals, would in a few thousand years stock the whole 
world. The increase of every species of monkey must be 
checked by some means; but not, as Brehm remarks, by the 
attacks of beasts of prey. No one will assume that the actual 
power of reproduction in the wild horses and cattle of America, 
was at first in any sensible d^iee incieased ; or that, aa each 
district became fully stocked, ttiis same power was diminished. 
No doubt in this case, and in all others, many checks concur, 
and different checks under different circnmstancea ; periodical 
dearths, depending on unfavourable seasons, being probably the 
most important of all. So it will have been with the early pro- 
genitors of man. 

NatuTui Sdection. — We have now seen that man is variable in 
body aud mind; and that the variations are induced, either 
directly or indirectly, by the samo general causes, and obey the 
some general laws, as with the lower animals. Man has spread 
widely over the fiice of the earth, and must have been exposed, 
during his incessant migrations,'* to the most diversified con- 
ditions. The inhabitauts of Tierra del Fuego, the Cape of Good 
Hope, and Tasmania in the one hemisphere, and of the Arctic 
regions in the other, must have passed through many climates, 

" tall of man. He shews Ihat the 

"by the many foul customs, es- 

" iastlncts of the higher animals 

" peoiaily as to marriage, of savage 

" are far nobler than the habits of 

'■ tribes. What does the Jewish 

" sarage raisa of men, and be finds 

" traditlM of the moral degeuera- 

" tion of man through hissuacehiug 

" re-introdnoe,— in a form of the 

" at a knowledge forbidden him 

" s'lhslantial jtthodoxy of whicli he 

" by his highest instinct assert 

"beyond this r 

" aai to introdnce as a EcientiSe 

« See some good remarks to thi< 

" hypothesis the docttine that man's 

(iTett by W. Stanley Jevons, " A 

" ;>un of iBOicfcd/. was th« cause of 

" deduction fiwn Darwin's Theorj,' 

" B temporary but long-endming 

*Natui-e,'i889, p. aSL 

'' moral deteiioratiou, us iuJicaCed 


4S The Descent of Man. Vx'xi L 

and changed their habits many times, lioforo they reached their 
present homes." The early progenitors of man must also Lavo 
tended, hku all other animals, to have increased beyond their 
meaaa of suhsiHtenee ; they must, therefore, ocoasicnally have 
been exposed to a stru^ie for esistenoc, and consequently to the 
rigid law of natural selection. Benoficial variutiocs of all kinds 
will thus, either occasionally or habitually, hare been preserved 
and injurious ones ehminated. I do not I'efer to strongly-marked 
deviations of structure, which occur only at long intervale of 
time, but to mere individual differences. Wo know, for instance, 
that the muscles of our handa and feet, which determine our 
powers of movement, are liable, like those of the lower animals," 
to ineessstnt variahility. If then the progenitors of maninhabit- 
ing any district, especially one undergoing some change in its 
conditions, were divided into two equal bodies, the one half 
which included all the individuals best adapted by their powers 
of movement for gaining subsistence, or for defending themselves, 
would on an average survive in greater numbers, and procreate 
more offspring then the other and less well endowed half. 

Man in the rudest state in which he now exists is the most 
dominant animal that has ever appeared on this earth. He has 
spread more widely that any other highly organised form : and 
all others have yielded before him. He manifestly owes this 
immense superiority to his intellectual faculties, to his social 
habits, which lead him to aid and defend his fellows, and to his 
corporeal structure.- The supreme importance of these characters 
has been proved by the final arbitrament of the tiattle for hfe. 
Through his powers of intellect, articulate language has been 
evolved ; and on this his wonderful advancement has mainly 
depended. As Mr. Chauncey Wright remarks ;'* " a psycholc^cal 
" analysis of the faculty of language shews, that even the smallest 
" proficiency in it might require more brain power than the 
" greatest proficiency in any other direction." He has invented 
and is able to use various weapons, tools traps &c with which 
he defends himself, tills or catches prey and otherwise ob aii.s 
food. He has made rafts or canoes for hshiag or crossmg ovtr 
to neighbouring fertile islands. He has discovered the art of 

IB Migra- " ciasaej id any t 

theit 'Anatomy of the 
(' Trausact. i^log, S 


caip. rr. 

Manner of Development. 


raftking fiio, br which hard and stringy roofs can bo rendered 
digestible, and poisOBous roots or herbs innoenonH, This die- 
cOTory of fire, probably tho greateet ever made b j man, escoptinj; 
language, dates from bofore the dawn of.historj. These severai 
inTontions, by which man in tto rndest state has become so pre- 
eminent, are the d jeet results of tho deTelopment of his powers 
of oiffiervation, memory, curiosity, ima^natJon, and reason. I 
cannot, therefore, understand how it is that Mr. Wallace" main- 
taiiK, that "natural selection could only have endowed tho , 
" savage with a brain a little superior to that of an ape." 

Although the intelleotnal powers and social habits of man arc 
of paramount importance to him, we must not uaderrata the 
importanae of his bodily structure, to which subject the remain- 
der of this chapter will be devoted ; the development of the in- 
tellectual and social or moral facnlljes being discnsscd in a later 

Even to hammer with precision is no easy matter, as every 
one who has tried to )eam carpentry will admit. To throw a 
stone with as true on aim as a Fcegian in defending himself, or 
in tilling birds, reqnires the most consummate perfection in tho 
correlated action of the mnseies of tlie hand, arm, and shoulder, 
and, further, a fine sense of touch. In throwing a stone or spear, 
and in many other actions, a man must stand firmly on his feet ; 
and this again demands the perfect co-adaptation of numerous 
muscles. To chip a flint into the rndest too!, or to form a 
(lorhed spear or hook from a bone, dtmands the use of a perfect 
hand; for, as a most capable Judge, Mr. Schoolcraft,'" remarlrs, 
the shaping fragments of stone into knives, Jancea, or arrow-heads, 

"''Quarterly Reriqw,' April 
I8S9, p. 3S2. This sabj«ct is more 
fully discusHd la Mr. VTalUce's 
'Contributions to the Thcocy of 
Hatmal Selection,' 1ST0, in which 
all the assays referrnl lo in this 
w k p bl hed r r 

Man has bee biy cised b) 

t qnoti 

t JUS 


htq Urn rselle, J 


Th mark q 
w 11 sn pn J 


ead M Wll 


Th Orlg 

t H m 


1 S 1 t Ei 

lly p h 

lished 1 b Ab h p 1 

gical R 

Tl w 

M y I8b+ p 1 i 

ramark by Sir 

J. Lubbock ('Pro- 

hiGtorio Times,' 

1865, p. 478) in 

paper, namely, that 

Ml. Wallace, " 

with choTBcteristic 

" nnselli^hiiess 

sscribea it (i e the 

deo f t 

lid!) 0- 

rvedlyt M 


1 h»d t b 

gh t wth th 

sam 1 bo 

t n, t tt m 

Q I d by M lawsu T 

h Law f N 

ral Sel t — 

I b1 Q t 

]yl ».l fMd 


50 "Ike Descent cf Man. Part 1 

shewB " extraordinary ability and long practiM." This ia to a 
great extent proved by tie fact that primeval men practised ft 
division of labour; each man did not majnifacture liis avm flint 
tools or rude pottery, but certain individuaLs appear to bave 
devoted themselves to snch work, no doubt receiving in exchange 
the produce of the chase. Archsologisfe are convinced that aa 
enormous interval of time elapsed before our ancestors thought 
of grinding chipped flints into smooth tools. One can hardly 
donbt, that a man-like animal who possessed a hand and arm 
suffloientlj perfect to throw a stone with precision, or to form a 
flint into a mde tool, could, with sufGeient practice, as far as 
mechanical skill alone ia concerned, mate almost anything 
which a civilised man can maka The structure of the hand in 
this respect may be compared with that of the vocal oi^ns, 
which in the apes are used for uttering various signal-cries, or, 
as in one genus, ronsicitl cadences ; but in man the closely 
similar vocal organs have becomb adapted through the inherited 
effects of use for the utterance of articulate language. 
/ Turning now to the nearest allies of men, and therefore to Iha 
beet representatives of our early progenitor, we find that the 
hands of the Quadrumana are constructed on the same general 
pattern as our own, but are far less perfectly adapted for diver- 
wfied uses. Their hands do not serve for locomotion so well 
as the feut of a dt^ ] as may be seen in such monkeys as the 
chimpanzee and orang, which walk on the outer margins of 
the palms, or on the knuckles." Their hands, however, are 
admirably adapted for climbing trees. Monkeys seize thin 
branches or ropes, with the thumb on one side and the fingert 
and palm on the other, in the same manner as we do. They can 
thus also lift rather large objects, such as the neck of a bottle, to 
their mouths. Baboons turn over stones, and scratch up roots 
with their hands. They seize nuts, insects, or other siuall 
objects with the thumb in opposition to the fingers, and no 
doubt they thus extract eggs and the young from the nests of 
birds. American monkeys beat thewildoranges on the branches 
until the rind is cracked, and then t«ar it off with the fingore of 
the two hands. In a wild state they break open hard frnit.s 
with stones. Other monkeys open mussel-shells with the two 
thumlo. With their fingers they pull out thorns and burs, and 
hunt for each other's paraeites. They roll down stones, or throw 
them at their enemies : nevertheless, they are clumsy in these 
various actions, and, as I have myself seen, are quite unaWe to 
throw a stone with precision. 

" Ob en, ' ADatomy of Vertebrates,' rol. i:i. p. 71. 


CiiAT. U. Manner of Dtvelopmmt. 51 

It seems to me far from true that becanse " objects are grasped 
" clmnsily" by monkeys, "a maeh. less specialised oi^an oi 
" prebensioa " ■would have served them " equallj well with 
tiieir present hanijs. On the contrary, I see no reason to doubt 
that more perfectly constructed hands would have been an 
advantage to them, provided that they were not thus rendered 
less fitted for chmbing trees. We may siapeet that o band as 
perfect as that of man would have been disadvantageous for 
climbing ; for the most arboreal monkeys in the world, namely, 
Atelee in America, CoJobus in Africa, and Hylobates in Asia, 
are either thumbless, or their toes partially cohere, so tbat their 
limbs are converted into more grasping hooks.'" 

As soon as some ancient member in the great series of the 
Primates came to be Iess arboreal, owing to a change in its 
manner of procuring subsistence, or to some cbange in the 
surrounding conditions, its habitual manner of progression would 
have been modified : and thus it would have been rendered more 
stiictly quadrupedal or bipedal. Baboons frequent hilly and 
rocky distriots, and only from necessity climb Mgh trees ; '' and 
tliey have acqniied almost the gait of a dog. Man alone has 
become a biped ; and we con, I think, partly see how ho has 
come to assume his erect attitude, which forms one of his most 
conspicuous characters. Man could not have attained his present 
dominant position in the world without the use of bis hands, 
which are so adndrably adapted to act in obedience to his will. 
Sir C. Bell '' insists tbat " the hand supplies all instruments, 
■' and by its correspondence with the intellect gives him univcr- 
" sal dominion." But the hands and arms could hardly have 
become perfect enough to have manufactured weapons, or to 
have hurled stones and spears with a tnie aim, as long as they 
were habitually used for locomotion and for supporting the 
whole we^ht of the body, or, as before remarked, so long as they 
were especially fitted for climbing trees. Such rough treatment 
would also have blunted the sense of toueb, on whieb tbeir 
delicate use largely depends. From these causes alone it would 
have been an advantage to man to become a biped; but for 

' QuiTterly Review,' Apvil 

tHo feel 

regulirly cohere ; aad this, as Mr. arboreal animals 

Blyth informs me, la oceasionallj wonderfutlj hook- 
tbe case with the toes uf H. agilis, " Brelim, 'Tb 

fcr, and te»ciscH3. Colobns is strictly 80, 
Mboreal and eitraordiaanly active " "The Hand. 

(BrchH, ' ThierUben,* B. i. t. SOX water Treatise,' 1) 


52 The Descent of Man. Paet l 

many actions it is indispensable that lie arms and whole uppei 
part of the body should be free ; and he must for this end stncd 
firmly on hia feet. To gain this great advantage, the feet ha^e 
beon rendered flat ; and the great toe has been peculiarly modi- 
fied, though this has entailed the almost complete loss of its 
power of prehension. It accords with the principle of tho 
division of physiological labour, preToiling throughout the 
animal kingdom, that as the hands became perfected for pre- 
hension, the feet should have become perfected for support and 
locomotion. With some savages, however, the foot has not 
altogether lost its prehensile power, as shewn by their manner 
of climbing trees, and of using them in other ways." 

If it be an advantage to man to stand firmly on his feet and to 
have his hands and arms free, of which, from his pre-eminent 
success in the battle of life, there can be no doubt, then I can see 
no reason why it should not have been Bdvantageous to the 
progenitors of man to have become more and more erect or 
bipedal. They would thus have been better able to defend 
themselves with stones or clubs, to attack their prey, or other- 
Tvise to obtain food. The best built individuals wonJd in the 
long run have succeeded best, and have survived in laiger 
numbers. If the gorilla and a few allied forms had become 
cEtinot, it might have been argued, with great force and apparent 
truth, that an animal could not have been gradually converted 
from a quadruped into a biped, as all the individuals in an 
intermediate condition would have been miserably ill-fitted 
for progression. Eut we know (and this is well worthy ot 
reflection) that the anthropomorphous apes are now aetually ia 
an intermediate condition ; and do one doubts that they are c 
the whole well adapted for their conditions of hfa Thus the 
gorilla runs with a sidelong shambling gait, but more commonly 
pri^ressos by resting on its bent hands. The long-armed apes 
occasionally use their arms like crutches, swinging their bodies 
forward between them, and some kinds of Hylobates, without 
having been taught, can walk or ran upright with tolerable 
([nlckness ; yet they move awkwardly, and much less securely 
than man. We see, in short, in esisting monkeys a manner of 
progTOBsion intormediate be i ween that of a quadruped and a 

" IKcke! lias rb eicelleut dis- foot as a prtbeosile oi^n by mao ; 

bGcameabipedi'NatUrlichsSchop- of prognssioa of the taigher apes, to 

fuagsgeschithte,' 1868, s. 507. Dr. which I allude in the following 

BUchner (' Conftrences ■nrla ITieorie paragraph : see aUn OweoC AofltoiBj 

!>arwiiiieiine,' IS69, p, 135) has of Vertebrates,' toI. iii. p. Tl) ai| 

giren good cawi of to* HM of tb* this latter aubjecL 


Uhap. iI. Manner of Developttent. J 3 

biped ; but, as an unprq'iadiced judge '" insists, the anthropomor- 
phous apes approach in structure more neariy to the bipedal 
than to the quadrupedal t;pe. 

As the progenitors of man became more and more erect, witQ 
their hands and arms more and more modified for prehension 
and other purposes, with their feet and legs at tJie tiame time 
transformed for firm support and progression, endless other 
chains of structure would have become necessary. The pelvis 
would have to be broadened, the spine peculiarly curved, and the 
head fixed ia an altered positiott, all which changes have been 
attained by man. Prof. Sohaaff hausen '° maintains that "the 
" powerful mastoid processes of the human skull are the result of 
" his erect position ;" and these prooessea are abscat in the oraug, 
chimpanzee, &c., and are smaller in tbo gorilla than in man. 
Various other structures, which appear connected with man's 
erect position, might here lave been added. It is very difficult 
to decide how far these correlated modifications are the result of 
natural selection, and how far of the inherited effects of the 
increased use of certain parts, or of the action of one part on 
another. No doubt these meausof change often co-operate ; thus 
when certain muscles, and the crests of bone to which they are 
attached, become enlarged by habitual use, this shews that 
certain actions are habitually performed and most be seiviceable. 
Hence the individuals which performed them best, would tend 
to survive in greater numbers. ^ 

The free use of the ajrms and hands, partly the cause aud \ 
partly the result of man's erect position, appears lo have led in an I 
indirect manner to other modifications of structure. The eai^^;_/ 
male forefathers of maa were, as previously stated, probably I 
furnished with great canine teeth; but as they gradually \ 
acquired the habit of using stones, clubs, or other weapons, for 1 
fighting with their enemies or rivals, they would use their jawa 
and teeth less and less. In this case, the jaws, together with the ( 
teeth, would become reduced in size, as we may feel almost sure 
from inmimerable analogous cases. In a future chapter we 
shall meet with a closely parallel case, in the reduction or com- 
plete disappearance of the canine teeth in male ruminants, 
apparently in relation with the development of their horns ; and 
in horses, in relation to their habit of fighting with their incisor 
teeth and hoofs. 

" Prof. Broea, La Coustitntioi tiia Skull,' tTaaslaled iu ' Anthro- 

aea Vei-tibrefl caudates, *La Revus pological Review,* Oct. 1868, p, 

d'Aathropologie,' 1871, p. 2S, 428. Owen (' Aoitomy of Verte- 

(Mpirata copy). inates,' vol. ii. 1866, p. 551) on the 

'*'» Pcimitivs Form of nmstoid processes in the higher ape* 


54 TJie Descent of Matt. 1'akt I. 

In the adult mde anthropomorphous apes, as Eiitimojer," 
ftud others, have insisted, it is the effect on the skull of the great 
development of tlie Jaw-mnaelea that causes it to differ so greatly 
in many respects from that of man, and has given to these 
animals "a ti'ulj frightful phjaiognomy." Therefore, as the jaws 
and teeth in man's progenitors gradually becamareducedin size, 
the adult skull would have come to resemble moro and mora 
that of existing man. As we shall hereafter see, a great reduction 
of the canine teeth in the males would almost certainly affect the 
teeth of the females through inheritance. 

As thevarious mental faculties gradually developed themselves 
the brain would almost certaijiiy become lai^er. No one, I 
presiune, donbts that the large proportion which the size of 
man's brain bears to his bodj, compared to the same proportioii 
in the gorilla or orang, is closely connected with his higher 
mental powers. We meet with closely anal<^ous facts with 
insects, for in ants the cerebral ganglia are of estraordinary 
dimeneionB, and in all the Hymenoptera these ganglia are many 
times larger than in the less intelligent orders, such »s beetles." 
j On the other band, no one supposes that the intellect of any 
\ two animals or of any two men can be accurately ganged by the 
' cubic contents of their skulls. It is certain that there may b« 
extraordinary mental activity with an extremely small absolute 
mass of nervous matter: thus the wonderfully diversified 
instincts, mental powers, and affections of ants are notorious,, 
yet their cerebral gangha are not so large as the quarter of a 
small pin's head. Under this' point of view, the brail 
one of the most marvellous atoms of matter in the wi 
more so than the brain of a man. 

The belief that there exists in man some close relation bi 
the size of the brain and the development of the inlellectiial 
faculties is supported by the comparison of the skulls of savage 
and civilised races, of ancient and modern people, and by the 
analogy of the whole vertebrate series. Dr. J. Barnard Davis has 
proved,™ by many careful measurements, that the mean internal 
capacity of the skull in Europeans is 923 cubic inches ; in 
Americans 87-5; in AsiatJCB 87-1; and in Australians only 81 '9 
cubic inches. Professor Broca^foimdthatthoniQeteenthcentnrj 

" 'Die Grenien der Thierwelt, vomileria,' 1870, p. 14. Mr son, 
eineBetriditangzu Darwin's Unre,' Mr, F. Darwin, disseeted for me the 
1868, ». 51. cerebral ganglia of the Fvnnica 

1850, p. 203. See also Mr. Lowne, 1869, p. 513, 
Anatomy and Phja. of the Musc/i " ' l.ea Stilectioas, SI. P, Eri-a, 


CHir. II. 

Manner of Development. 


Bkolla from graves in Paris were larger than those bom TBolta 
of the twelfth century, in the proportion of 1184 to 1423; and 
that tho increased size, as ascertained b; measurements, was 
esclnsiTelj in the frontal part of the skull — the seat of tho 
intellect\ial faculties. Priohard is persuaded that tho present 
inhabitants of Britain have " much more capacious brain-cases " 
than the aoaoient inhabitants. Neverthelesa, it must be admitted 
that some skullH of very high antiquity, such as the famous ono 
of Neanderthal, are well developed and capacious." With 
respect to the lower animals, M. E. Lajtet,*^ by comparing the 
crania of tertiary and recent jnammals belonging lo tho same 
groups, hna come to the remarkable conclusion that the brain is 
generally larger and the convolutions are more complex in the 
more recent forms. On the otter hand, I have shewn" that the 
brains of domeistjc rabbits are considerably reduced in hulk, in 
comparisonwith those of thewild rabbit or hare; and this may be 
attributed to their having been closely confined during many 
generations, so that they have exerted their intellect, instincts, 
flenses and voluntary movements but little. 

The gradually increasing weight of the brain and skull ini 
man must have influenced the development of the supporting 
spinal column, more especially whilst he was becoming erect l 
As this change of position was being brought about, the internal ' 
pressure of the brain will also have in9nenoed the form of the 
skull; for many facts show how easily the skull is thus affected. 
Ethnologists believe that it is modified by tlie kind of cradle in ''. 
which infants sleep. Habitual sjMisms of tho muscles, and a 
eieatrix from a severe burn, have permanently modified the facial 
bones. In youi^ persons whose heads have become fixed either 
Bideways or backwards, owing to diseiee, one of tho two eyes has 
changed its position, and the shape of the skull has been altoted 

Revue d'Anthropologies,' 1873; 

tbe other hand, with aavages, th 

i^e also, as quoted in G Vogfa 

rage inelodes only tbe more ca 

' Leotures on Man,' Eng. tranalat. 

individuals, who have Iwen a 

1864, pp. 88, 90. Prictard, ' Php. 

Burviva onder eitremely hard 

Hist, of Mankind,' vol. i- 1338, p. 

ditioui of life. Broca thus ei 


the otherwiao ineiplicable fac 

•1 In the interesting article just 

th« oieau capacity of the at 

(oietrod to, Prof. Broca has well 

the ancient Troglodytes of Lo 

remarked, that iu civilised nations, 

greater than that of modern F 

the average capacity of the skull 

must 1* lowered by the preserva- 
tion of a»^lB number of 

" ' Comples-rendus des Sd- 

&c June 1, 1868. 

" 'The Variation of Animal 

s, the ave- 

■mptly Plnnla under /'-meat 


S6 The Descmt of Man. I'iai 1. 

spparently by the pressure of the brain in a new diraotioii."' 
IhttYoshewn that with long-eared rabbits even Wi ttiftmga cauae 
as the lopping forward of one ear drags forward ahnost everj 

ft)one of the skull on that side ; so that the bonca on the opposite 
Bide no longer strictly coirespoiid. Lastly, if any animal were 
to increase or diminish, much in general size, mthout any change 
in its mental powers, or if the mental powers were to bo much 
inoreaaed or diminished, without any great change in the sizo of 
the body, the shape of the skull would almost certainly be 
altered. I infer this from my observations ou domestic rabbits, 
some kinds of which have become very mucli larger than the 
wild animal, whilst others have retained nearly the same size, 
but in both eases the brain has been much reduced relatively to 
the size of the body. Now I was at first much surprised on 
Snding that in all these rabbits the skull had become elongated 
or dolichocephalic; for instance, of two skulls of nearly eciual 
breadth, the one from a wild rabbit and the other from a large 
domestic kind, the former was 315 and flie latter 4'3 inches in 
length.* One of the most marked distinctions in different races 
of men is that the skull in some is elongated, and in others 
rounded; and here the explanation su^ested by the case of the 
rabbits may hold good ; for Welcfcer finds that short " men inchiie 
more " to brachyeephaly, and tall men to dolichocephaly ;"** aud 
tall men may be compared with the larger and longer-bodied 
rabbits, all of which have elongated skulls, or ax'e dolicho- 

From these several facts we can understand, to a certain 
extent, the means by which the great size and more or loss 
rounded form of the skull have been acquired by man ; and these 
are characters eminently distinctive of him in comparison with 
the lower animals. 

Another most conspicuous difference between man and the 
lower animals is the nakedness of his skin. Whales and 
porpoises (Cetacea), dugongs (Sirenia) and the hippopotamus ara 
naked; ajjd this may be advantageous to them for gliding 

" Schaaffhausen gives fi-om Blu- m 
luenljach and Busoh, the eases of the ht 

' the modification of the skull fr 

eition. He believes th&C ia c 
in trades, sacii m Uiut of u sh 

iliar. where 

tl.e head >s habil 


Id iirward 

>rehead be> 

ire rouQded 

1 and p 

" ' Variati 

on of 


■ &c.. 

L i. p. m 

e skall ; p. 

119, 1 

on the effect of 

slopping of one ei 

« Quo«d 

by Se 



. Revi. 

eiv,' Oct. 



Chap. II. Manner of Development 57 

tJmiugli the wat«r; sor would it bo iiijurioiis to them from tLo 
loss of warmth, as the species, which inhabit the colder i^ions, 
lite protected by a thick layer of blubber, eervit^ the Kama 
pm;pu£e as the fur of seals and otters. Elephants sod rhiuu' 
ceroses are almost hairless ; and as certain extinct species, 
which formerly lived under an Arctic climate, were covered with 
long wool or hair, it wotild almost appear as if the existing 
epooies of both genera had lost their hairy covering from exposure 
to heat. This appears the more probable, as the elephants in 
India which live on elevated and cool districts are more hairy "' 
than those on the lowlands. May we then infer that roanN 
became divested of bair from having aboriginaliy inhabited some 1 
tiMpieal land ? That the hair is chiefly retained in the male sex on ) 
the chest and face, and in both sexes at the junction of all four 
limbs with the tmnt, favours this inference— on the assumption 
that the hair was lost before man became erect ; for the parts 
which now retain moat hair would then hate been most protected 
from the heat of the sun. The crown of the head, however, 
ofiers a curious exception, for at all times it must have been one 
of the most exposed parts, yet it is thickly clothed with hair. 
The fa«t, however, that the other members of the order of 
Primates, to which man belongs, although inhabiting various hot 
regions, are well clothed with hair, generally thickest on the upper 
surface,"* is opposed to the supposition that man became nakiid 
through the action of the sun. Mr. Belt believes " that within 
the tropics it is an advantage to man to be destitute of hair, as 
he is thus enabled to free himeelf of the multitude of ticks (aoari) 
and other parasites, with which he is often infested, and which 
sometimes cause ulceration. But whether this evil is of sufficient 
magnitude to have led to the denudation of hia body through 
natural selection, may be doubted, since none of the many 
quadiTjpeds inhabiting the tropics have, as far as I know, 
acquired any specialised means of relief. The view which st 

to mo the most probable is that : 

man, or rather primarily woman, 

" Owen, 'Aoatoiur of Verl*- 

ever, states that in tbe Gorilla tbe 

OTAtes,' vqI. iii. p. 610. 

h^uf ia tliinner an the back, «hei» 

" Isidore Geoffrov St.-Hilaire re- 
marks (' Hist. Nst. GinSrale,' torn. 

it is partly rubbed off, thsn on tbe 
lower sur&ce. 

li 1859, pp. 2X5-217) on the head of 

•• TliB ' Naturalist in Nicaragua,' 

man being covei'ed with loag hair ; 

1874, p. 209. As some oonfitma- 

also on tha upper sortaeej of mon- 

tion of Mr. Bolt's view, J may quott 

keys and of other maromais being 

the following passage from Sir W. 

mote tbickly clothed than the lower 

Denuon ('Varieties of Vioe-Begal 

surfaces. This has likewise l>een 

life,' ToL i. 1870, p. 440): " It b said 

Diiserved by varioiu authors. Pi'ot 

"CO be a practice with the Aus- 

P. Ger-ais (' Ulst. Nat. des Mam- 

" tralians, when the venuio gel 

miftraH,' lorn. <. 1654, p. 38), bow- 


S$ The Descent of Mat. Vxm I. 

bocamo divested of hair for ornamental purposes, as we shall set. 
under Sexual Selection ; and, according to this belief, it is not 
Eurprising that man should differ so greatly in hairiness &om al ! 
other Primates, foi characters, gained through sexual seleotiou, 
often differ to an estxaordinary degree in closely-related forms. 

! According to a popular impression, the absence of a tail is 
utnincatly distinctiTe of man ; but as those apes which coma 
nearest to him arc dostitute of this oi^n, its disappearance does 
not relate exclusively to man. The tail often differs remarkably 
in length, within the same genus ; thus in some species of Ma«ao«s 
it is longer than the whole body, and is formed of twenty-four 
vertebrje; in others it consists of a scarcely visible sttimp, 
containing only three or foitr vertebrie. In some kinds of 
baboons there are twenty-five, whilst in the raandrill there are 
ten very small stunted caudal vortebrffi, or, according to Ouvier," 
sometimes only five. The tail, whether it be long or short, almost 
always tapers towards the end ; and this, I presume, results from 
the atrophy of the terminal muscles, together with their arteries 
and nerves, throi^h disuse, leading to the atrophy of the terminal 
bones. But no oiplanation can at present be given of the great 
diversity which often occurs in its length. Here, however, we 
are more specially concerned with the complete external dis- 
appearance of the tail. Professor Brooa has recently shewn" 
that the tail in all quadrupeds consistB of two portions, gfflierally 
separated abruptly from each other; the basal portion consists 
of vertebrje, more or less perfectly ehaimeUed and furnished with 
apophyses like ordinary vertebrse ; whereas those of the terminal 
portion are not channelled, are almost smooth, and Bcarcely 
resemble true vertebrte. A tail, though not externally visible, is 
really present in man and the anthropomorphous apes, and is 
constructed on exactly the same pattern in both. In the terminal 
portion the vertebrte, constituting the os coccya:, are quite 
rudimentary, being much reduced ia size and number. In the 
basal portion, the vertebrte are likewise few, are united fii'mly 
tegether, and are arrested in development; but they have been 
\ rendered much broader and flatter than the corresponding 
\ vertebra in the fails of other animals ; they constitute what 
Broca calls the accessory sacral vortebrse. These are of functional 
importance by supporting certain internal parts and in other 
ways ; and their modification is directly connected with tho erect 

M). St. George Mivart, 'Pros. Geoffroy, 'Hist. Sat. Gto.' lom. ii. 

l>r. JV 

'e. gWj, 'Ca 
;«uins.' Owen, 
itebratea,' vol :;. j 

Tie d'AathropoIogie,' 1878; 
titutioD ias Vei-t*brsi can- 


Oiui'. II. Manner of Development. 59 

or semi-erect attitude of man and the anthropomorphona apee. 
This conclusion is the more tmatwotthy, as Bioca formerly held 
a different Tiew, which he has now ahandoned. The modifica- 
tion, therefore, of the liaaal caudal yertebrte in man and the 
higher apes may have been effected, directly or indirectly, 
through natTU'sl selection. 

But what are we to aay about the rudimentary and variable 
Tertebrte of the terminal portion of the tail, forming the os coixyT. ? 
A notion which has often been, and will no doubt again be 
ridiculed, namely, that friction baa had something to do with 
the disappearance of the external portion of the tail, is not 
eo ridiculous as it at first appears. Dr. Anderson " states 
that the extremely short tail of Macacus brunneus is formed of 
elovea vert«br!B, inclndii^ the imbedded basal ones. The- 
extremity is tendinous and contains no vertebrte; this is suc- 
ceeded by five rudimentary ones, so minute that together they 
are only one line and a half in length, and these are permanently 
bent to one side in the shape of o hook. The free part of tha 
tail, only a Uttlo above an inch in length, includes only four move 
small Tertebrie. This short tail is carried erect ; but about a 
quarter of its total length is doubled on to itself to the left; and 
this terminal part, which includes the hook-liie portion, serves 
" to fill up the interspace between the upper divei^nt portion 
" of the callosities ;" so that the animal sits on it, and thus renders 
it rough and callous. Dr. Audersoa thus suios up Lis observa- 
tions: "These facts seem to me to ha™ only one explanation; 
" this tail, from its short size, is in the monkey's way when it 
" sits down, and frequently becomes placed under the animal 
" while it is in tins attitude ; and from the circumstance tliat it 
" does not estend beyond the extremity of the ischial tuberosities 
" it seems as if tlie tail originally had been bent round, by the 
" will of the animal, into the interspace between the callosities, to 
" escape being pressed between them and the ground, and that 
" in time the curvature became permanent, fitting in of itself 
" when the oi^n happens to be sat, upon." Under these circum- 
stances it is not surprising that the sur^ice of the tail should 
have been roi^hened and rendered callous ' and Dr. Murie,'™ who 
carefully observed this species in the Zoolc^cal Gardens, as well 
BS throe other closely allied forms with shghtly longer tails, says 
that when the animal eits down, the t&H " is necessarily thmst 
" to one side of the buttocks ; and whether long or short its root 
■ is consequently liable to be rubbed or chafed." 


t. Zuo.og. 3oc.,' 1872, p. •» ' ProD. Zoolog Soe. 1872, p 



6o Tfu Descejit of Man. Vaoi I 

h»ve evidence that mutilations occasioaally produce ai! inhoritod 
effect," it is notveryimprobable tliat inshortrtailedmoiitejB.the 
projecting part of tlie tail, being functionally useless, should after 
many generations have become rudimentary and distorted, from 
being COTitinually rubbed and chafed. We see the projecting part in 
this condition in the Macacits brunneas, and absolutely abortud in 
th.6 M.eca.iuiatai and in several of the higher apes. Finally, Uien, 
as Ikr a^ we can judge, the tail has disappeared in man and the 
anthiopomorphoas apes, owing to the terminal portion having 
been injured by friction during a long lapse of time ; the basal 
and embedded portion having been reduced and modified, so as 
to become suitable to the erect or semi-erect position. 

I have now endeavoured to shew that some of the most 
distinctive characters of man have in all probability been 
acquired, either directly, or more commonly indirectly, tlirougb 
natural selection. We should bear in mind that modifications 
in strncture or oonsfitution, which do not serve to adapt an 
organian to its habits of life, to the food which it consumes, or 
passively to the surrounding conditions, cannot have been thus 
acquired We must not, however, be too confident in deciding 
what modifications are of service to each being : we should 
remember how httle we know about the use of many parts, or 
what changes in the blood or tissues may servo to fit an 
organism for a new climate or new kinds of food. Nor must we 
foiget the principle of correlation, by which, as Isidore Geoffroy 
has shewn in the case of man, many Btrange deviations of 
etructuie are tied together. Independently of correlation, a 
change in one part often leads, tlirougli the increased or decreased 
use of other parts, to other changes of a quite unexpected 
nature. It is also well to reflect on such facts, as the wonderful 
growth of galls on plants caused by the poison of an insect, and 
on the remarkable changes of coloiu' in the plumage of parrots 
when fed on certain fishes, or inoculated with the poison of 
toads ;'^ for we can thus see that the fluids of the system, if 
altered for some special purpose, might induce other changes. 
We shonld especially boar in mind that modifications acquired 

"'s inherited effects of mot-inots liitiug 
observations on the transmitted off the barba of their owa tail. 

effect of hh operation causing epi- 

feathers. See also on the general 

lepsy in gninaa-pigs, and likewise 

subject 'Variation of Animals and 

more receotl; ou the Hualogous 

Piabts undei' Domer^ticution,' vol. 

eHeets of cutting tlie sympathetic 

ii., pp. 22-24. 

nerve in the neck. I shall hereafter 

" ' Tlie Variation of Animal, md 

have occasion to relei- to Mr. Salvin., 

Plants under Donaestieaticn,' vol. ^ 


Onip. IL Manner of Development. <*^ 

and continnally used during past ages for gome nsoful purpose, 
would probably become firmly fised, and might bo Jong jnliejited. 

TliTis a large yet nndefln&i extension may safely be giyen to 
the direct and indirect results of natural selection; but I now 
admit, after reading the essay by Niigeli on plaiite, and the 
remarks by Tarioua authors wiUi respect to animals, more 
especially those recently made by Professor Broca, that in the 
earlier editions of my ' Origin of Species ' I perhaps attributed 
too much to the action of natural selection or the emrrival ol 
the fittest. I have altered the fifth edition of the ' Origin ' so as to 
confine my remarks to adaptive changes of stracture ; but I am 
conTinoed, from tho light gained during even the last few years 
that very many structures which now appear to us useless, will 
hereafter he proved to be useful, and wil! therefore come within 
the Ho^e of natural eeleetion. Nevertheless, I did not formerly 
consider sufficiently the existence of structures, which, as far aa 
we can at present judge, are neither tieneficial nor injurious 
and this I believe to be one of the greatest oversights as yet 
detected in my work. I may be permitted to say, as some 
excuse, that I had two distinct objects in view ; firstly, to 
shew tbat species had not been separately created, and seeondiy, 
that natural selection hsd been the chief agent of change, 
though largely aided by the inherited effects of habit, and slightly 
by the direct action of the surroundiag conditions. I was 
not, however, able to annni the influence of my formet belief, 
then almost uniyersftl, that each speeiiis had been purposely 
created ; and this led to my tacit assumption that every detail 
of structure, excepting rudiments, was of some special, though 
unrecc^msed, service. Any one with this assumption in his 
mind would naturally extend too for the action of natural 
selection, either durii^ past or present times. Some of those 
who admit the principle of evolution, but reject natural Beleo- 
tion, seem to foj^t, when criticising my book, that I had the 
above two objects in -view ; hence if I have erred in giving to 
natural selection great power, which I am very far from 
ftdraittii^, or in having es^^erated its power, which is in itBOlf 
probable, I have at least, as I hope, done good service in aiding 
to overthrow the d(^ma of separate creations. 

It is, as I can now see, probable that all organic beings, 
including man, possess peculiaritiea of structure, which nei13ier 
are now, nor wore formerly of any service to them, and which, 
therefore, are of no physiological importance. We know not 
what produces the numberless slight differences between tho 
individuals of each species, for reversion only carries the 
prohloni a few steps backwards; but each peculiarity mijst 


62 Tfie Descent of Man. Part I 

hftve had ite efEciect caiise. If these canses, mhatofer thej 
may be, wore to act more uniformlj and energetically during a 
lengthened period (and against this no reason eaji be assigned), 
the result woald probalilj be not a mere slight individual 
, difference, but a well-marked and constant modificatjou, though 
! one of no physiological importance. Chwiged etrnctures, which 
are in no way beneficial, cannot be kept uniform througii natural 
■, selection, thongli the injurious will be thus eliminated. Uni- 
formity of character would, however, naturally follow from the 
assumed uniformity of the exciting causes, and likewise from 
the free intercrossing of many icdivi duals. During successive 
periods, the same organism might in this manner acquire 
successive modificatiouB, which would he transmitted in a nearly 
uniform state as long as the eiciting causes remained the same 
and there was free intercrossing. With respect to the exciting 
causes we can only say, as wlaen speaMng of so-called spon- 
taneous Tariations, that they relate much more closely to the 
constitution of the varying oiganism, than to the nature of the 
conditions to wliich it has been si ' ' 

Conclusion. — In tiiis cliapter we have seen that as man at the 
present day is liable, like every other animal, to multiform 
individual differences or shght variations, so no doubt were the 
early pri^enitors of man ; the variations being formerly induced 
by Vae same general causes, and governed by the same general 
^^nd complex laws as at present. As all animals tend to multiply ■ 
beyond their means of subsistence, so it must have been with 
the progenitors of man; and this would ineyibibly lead to a 
struggle for existence and to natural selection, the latter 
process would be greatly aided by the inherited effects of the 
increased use of parts, and these two processee would incessantly 
react on each other. It appears, also, as we shall hereafter seo, 
that various nmimportant characters have been acquired by man 
through sexual selection. An unexplained residuum of change 
must be left to the assumed uniform action of those unknown 
^encies, which occasionally induce strongly marked and abrupt 
deviations of structure in our domestic productions. 

Judging from the habits of savages and of the greater number 
of the Quadrumana, primeval men, and even their ape-like 
prt^enitors, probably lived in soeiety. With strictly social 
animals, natural selection sometimes acts on the individual, 
through the preservation of variations which are beneScial to 
the commum'ty. A community which includes a large number 
of well-endowed individuals increases in number, and is victo- 
rious over other less favoured ones; even although each separate 



Oe»p. 11, Manner of Development. 63 

membei^ gains no advantage over the others of the saflie com- 
munity. AseocJat«d iiiBocts have thus acquired many remark- 
able atructnreB, which aro of httle or no service to the individnal, 
Bnch as the pollen -coUeotii^ apparatus, or the sting of the 
worker-bee, or the great jaws of soldier-anta. "With the higher 
Boeial animals, I am not aware that amy stracture has been 
modified aolelj for the good of the coramunity, though some are 
of aecondary service to it For instance, the horns of nuninanis 
and the great canine teeth of baboons appear to have been 
aw|uired by tho males as weapons for sexual strife, but they are 
used in. defence of the herd or troop. In regard to certain 
mental powers the case, as we shall see in the fifth chapter, is 
wholly different ; for these faculties have been chiefly, or even 
eiclusively, gained for the benefit of the corarQimity, and the 
indiridnalB thereof, have at the same time gained an advantage 

It has often been objected to such views as the foregoing, that 
man is one of the most helpless and defenceless creatures in the 
%vorld; and that during his early and less well-developed 
condition he would have been still more helpless. The Duke of 
Ai^ll, for instance, insists'* that "the human frame has 
" diverged torn the stmoture of brutes, in the direction of 
" greater physica! helplessness and weakness. That is to say, it 
" is a divergence which of all others it is most impossible to 
" ascribe to mere natural seloctjon." He adduces the naked and 
unprotected state of the body, the absence of great teeth or 
clows for defence, the small strength and speed of man, and his 
slight power of discovering food or of avoiding danger by smell. 
To these deficiencies there might be added one still more 
serious, namely, that he cannot climb quickly, and so escape 
from enemies. The loss of hair would not have been a gi^at 
injury to the inhabitants of a warm country. For we know that the 
unclothed Fuegians can exist under a wretched climate. When 
we compare the defenceless state of man with that of apes, wo 
must remember that the great canine teeth with which the latter 
are provided, are poBBessed in their full development by the males 
alone, and are chiefly used by them for fightii^ with their rivals; 
yet the females, which ore not thus provided, manage to surviva. 

In regard to boddy size or strength, we do not know whether 
man is descended from some small species, like the chimpanzee, 
or from one as powerful as the gorflla ; and, therefore, we cannot 
say whether man has become larger and stronger, or smaliOT 

•* 'Primeval Man,' ]8fi9, p. 66 


64 Tlie Descent of Man. \_^ I'jht !, 

and weaker, tlian his ancestors. We eIiouM, bowcTer, i beat in 
mind that an animal possessing great size, strength, and .ferocity, 
and which, hke the gorilla, could defend itself from all CnemieB, 
would not perhaps have become social; and this would most 
effectually have checked the acquirement of the higher^ mental 
qualities, such as eympatliy and the love of his fellows. Hence it 
might have been an immense advantage to man to have sprung 
from some comparatively weak creature. 

The small strength and speed of man, his want of natural 
■weapons, &,c~, are more than counteibalanced, firstly, by his 
intellectual powers, through which he has formed for himself 
■weapons, tools, &c., though still remaining in a barbarous state, 
and, secondly, by his Bodal qiiaJities which lead him to give and 
receive aid from his fellow-men. No oonntry in the world abonnds 
in a greater degree with dangerous beasts than Southern Africa; 
no country presents more fearful physical hardships than the 
Arctic regions; yet one of the puniest of races, that of the 
Bushmen, maintains itself in Southern Africa, as do the dwarfed 
Esquimatii in the Aretio regions. Theanoostors of man were, no 
doubt, inferior in intellect, and probably in social disposition, fo 
the lowest esisting savages ; but it is quite conceivable that they 
might have existed, or even flourished, if they had advanced iu 
intellect, whilst gradually losing their brnte-like powers, such 
as that of climbing trees, &.C. Biit these ancestors would not 
have been exposed to any special danger, even if far more 
helpless and defenceless thaii any esisting savages, had they 
inhabited some warm continent or large island, such as 
Anstralia, New Guinea, or Borneo, which is now the home of the 
orang. And natiiral selection arising from the competition of 
tribe with tribe, in some such lai^ area as one of these, together 
with the inherited effects of habit, would, under fiivonrable 
conditions, have sufficed to raise man to bis presCTit high position 
in tlio crga^ic scale. 


Mental Powers. 

CoMP4BtsoN OF -THB Mental Powbes of Mah a 

IiOWBB Aniuals. 

cnUl power between the highest 

—Belief in 

We have seen in the last two chapters that man hears in his 
bodily strncture clear traces of his descent from some lower 
form ; but it may be urged that, as man diffiers so greatly in 
hia mental power from all other animals, there must be some 
error in this conclusion. No douht the difference in this 
respect is enormous, even if we compare the mind of one of the 
lowest savages, who has no words to express any number higher 
than four, and who uses hardly any abstract terms for common 
objects or for the affections,' with that of the most highly 
organised ape. The difference would, no donht, still remain 
immense, even if one of the higher apes had been improved or 
ciTiIised as, much as adog has been in comparison with its 
parent-form, the wolf or jackal. The Fuegians rank amongst 
the lowest barbarians; but 1 was continually stmtk with 
surprise how closely the three natives on board H.M.S. " Beagle," 
who had lived some years in England, and could talk a little 
English, resembled us in disposition aad in most of our mental 
faculties. If no oi^nic being excepting man had possessed aay 
mental power, or if his powers had been of a wholly different 
natui'e.from those of the lower animals, then we should nevei 
have been able to convince ourselves that our high facnlties 
had been gradually developed. But it can be shewn that there 
is no fimdfljnental difference of this kind. Wo must also admit 
that there is a much wider interval in mental power between 
one of the lowest fishes, as a lamprey or iaiicelet, and one of the 
higher apes, than between an ape and man ; yet this interval 
is filled up by numberless gradations. 

Nor is the difference slight in moral disposition between a 
barbarian, such as the man described by the old navigator 


I by LubbMk, ' i'rehistorio 


66 The Descent of Man. 1'abt l, 

I Bjron, who dashed bis child aa the rocks for dropping a basket 
. of sea-iirohinB, and a Howard or Clarkson ; aod in infollect, 
; butween a savage who iises hardly any abstract terms, and a 
) Newtoa or Shakspoare, BifCetenees of this kind between the 
highest men of the highest races and the lowest savages, are 
1 connected by the finest gradations. Therefore it is possible that 
■ they might pass and be developed into each otiier. 
1 My object in this chapter is to shew that there is no fnnda- 
1 mental difference between man and the higher mammals in their 
1 mental faculties. Each division of the subject might have tieen 
extended into a separate essay, but must here be treated briefly. 
As no classiScation of the mental powers has been nniTersally 
accepted, I shail arrange my remarks in the order most con- 
venient for my purpose ; and will select those facts which have 
struck me most, with the hope that they may produce some 
effect on the reader. 

With respect to animals very low in the scale, I shall give 
some additional facts under Sexual Selection, shewing that their 
mental xwwers are much higher than might have been expected. 
The Tariability of the faeultiea in the individuals of the same 
species is an important point for us, and some few illustrations 
will here be given. But it would be superfluous to enter into 
many details on this head, tor I have found on freqnent enquiry, 
that it is the unanimous opinion of all those who have long 
attended to animals of many kinds, including birds, that the 
individuals differ greatly in every mental characteristic. In 
what manner the mental powers were first developed in the 
lowest oi^anisms, is as hopeless an enquiry as how life itself 
first originated. These are problems for the distant future, if 
they are ever to l>e solved by man. 

fundamental intuitions must be the same. Man has also some 
few instincts in common, as that of self-preservation, sexual love, 
the love of the mother for her new-born offepriag, the desire 
possessed by the latter to suck, and so forth. But man, perhaps, 
has somewhat fewer instincts than those possessed by the 
animals which come next to him in the series. The orang in 
r the Eastern islands, and the chimpanzee in Africa, build plat- 
I forma on which they sleep ; and, as both species follow the same 
\ habit, it might be ai^ed that this was due to instinct, but wo 
] cannot fei.1 sure that it is not the result of both animals having 
j similar want"?, and possessing similar powers of reasoning. 
These apes, as we may assume, avoid the many poisonoiis fruits 
of tie tropics, and man has no such knowledge ; but ofl our 
d<ime6tic animals, when taken to foreign lands, and whun first 


OiiAP, ni. Meitlal Powers. by 

turned out m tlio spring, often eat poisonous licrbs, which tliey 
afterwards avoid, we cannot feel sure that the apes do not learn 
from theii otto experience or from that of their parents what 
fruits to select. It is, however, certain, as we shall presently so«, 
that apes havo an instinctive dread of serpents, and probably t/i 
other dangerous animals. --^ 

The fewness and the comparativo simplicity of the instincts in 
the higher animals are remarkable in contrast with those of the 
lower animals. Cuvier maintained that instdnct and intelligence 
stand in an inverse ratio to each other ; and some have thonght 
that the intellectual faculties of the higher animals have been 
gradually developed from their instincts. But Fouchet, in an 
interesting essay,' has shewn that no such inverse ratio really 
exists. Those insects which possess the most wondeiful instincts 
are certainly the most intelligent. In the vertebrate series, the 
least intelligent members, namely fishes and amphibians, do not 
possess complex instincts ; and amongst mammals the animal 
most remarkable for its instincts, namely the beaver, is highly 
intelh'gent, as will be admitted by every one who has read Mr. 
Morgan's excellent work.' 

Although the first dawnings of intelligence, according to Mr. 
Herbert Spencer,' have been developed through the multipiiea- 
tion and co-ordination of reflex actions, and althongh many of 
the simpler instincts graduate into reflex actions, and can hardly 
be distinguished from them, as in the case of young animal)' 
sucking, yet the more complex instincts seem to have originatea 
independently of intelligence. I am, however, very far from 
wishing to deny that instinctive actions may lose their fixed and 
untaught character, and be replaced by others performed by the 
aid of the free will. On the other hand, some intelligent actions, 
after being performed during several generations, become con- 
verted into instincts and are inherited, as when birds on oceanic 
islands leom to avoid man. These actions may then be said 
to be degraded in character, for they are no longer performed 
through reason or from experience. But the greater number of 
the more complex instincts appear to have been gained in a 
wholly different manner, through the natural selection of varia- 
tions of simpler instinctive actions. Such variations appear to 
arise from the same unknown causes acting on tJie cerebral 
oiganisatioB, which induce slight variations or individual dif- 
ferences in other parts of the body; and these variations, owing 

■ ' The American Bearer and L:. 


OS The Deicent of Man. Past 1 

to our ignorance, are often said to arise spontaneously. We ran, 
I think, come to no otiier conclusion with respect to the origin of 
the mote complex instincts, when we reflect on the matvelloua 
instincts of sterile workei-ants and bees, which leave no off- 
Epiing to inherit the elfects of experience and of modified hahits. 

Although, as wo Icara from the ahoTe-mentioned insects and 
the heayer, a high degree of inteUigenee is cei'tainlj compatible 
with complex instiucts, and although ooUnns, at first learnt 
voluntarily can soon through habit he perfonned with the 
quickness and certainty of a reflex action, jet it is not improbable 
that there is a certain amount of interference between the 
development of free intelligence and of instinct, — which latter 
implies some inherited modification of the brain. Little is 
known about the functions of the brain, but we can perceive 
that as the intellectual powers become biglily developed, the 
various parts of the brain must be connected by very intricate 
channels of the freest intercommunication ; and as a conse- 
quence, each separate part would perhaps tend to be less well fitted 
to answer to particular sensations or associations in a definite 
and inherited — that is instinctive^ manner. There seems even 
to exist some relation between a low degree of intelligence and a 
strong tendency to the formation of fixed, though not ichorited 
habits ; for as ft sagacious physician remarked to me, persona 
who are slightly imbecile tend to act in everything by routine 
or habit ; and they are rendered much happier if this is en- 

I have thought this digression worth giving, because we may 
easily underrate the mental powers of the higher animals, and 
especially of man, when we compare their actions founded on the 
memory of past events, on foresight, reason, and imagination, 
with exactly similar actions instinctively performed by the lower 
animals ; in this latter case the capacity of performing such 
actions has been gained, step by step, through the variability of 
the mental organs and natural selection, without any o 
intelligence on the part of the animal during each s 
generation. No doubt, as Mr. Wallace has argued,' much of the 
intelligent work done by man is due to imitation and net to 
reason; but there is this great difference between his actions 
and many of those performed by the lower animals, namely, that 
man cannot, on his first trial, make, for instance, a stone hatchet 
or a canoe, through his power of imitation. He has to learn his 
work by practice ; a beaver, on the other hand, can make its 
dam or canal, and a bird its neet, as. well, or nearly as well, and 

» 'CuBiribulionatotheTheoty if HaturalSeltrtion,' 1870, f. SIX 

■— ' '— Hostedby VjOOQIC 

Chap. IIT. Menial Powers. 6g 

a spider its wonderful web, quite as well," the first time it tries 
as when old and esperienced. 
, To return to our immediaifl subject : fho lower animals, lika 
man, manifeBtly feel pleasure and pain, happiness and misery. 
Happiness is never better exhibited than by young animals, such 
as puppies, kittens, lambs, &c., when playing together, like onr 
own childreiL Even insects play together, as has been described 
by that excellent observer, P. Hnbei,' who saw ants chasing and 
pretending to bite each other, like so many puppies, 

Tho fact that the lower animals are excited by the Kanio 
emotions as ourseWes is so well established, that it will not ba 
necessary to weary the reader by many details. Terror acts in 
tho same manner on them as on us, causing the muscles to 
tremble, the heart to palpitate, tho sphincters to bo relaxed, and 
the hair to stand on end. Suspicion, the offspring of fear, is 
eminently characteristic of most wild animals. It is, I think, 
impossible to read the account given by Sir E, Tennent, of the 
behaviour of the female elephants, used as decoys, without 
admitting that they intentionally practise deceit, and well know 
what they are about Courage and timidiiy are estremely 
variable qualities in tho individuals of the same species, as is 
plainly seen in ourdi^. Some dogs and horses are ill-tempered, 
and easily turn sulky; others aro good-tempered; and these 
qualities are certainly inherited. Every one knows how liable 
animals are to furious rage, and how plainly they show it. 
Many, and probably true, anecdotes have been published on the 
long-delayed and artful revenge of various animals. The 
accurate Bcngger, and Brehm ' state that the American and 
African monkeys which they kept tame, certainly revenged 
themselves. Sir Andrew Smith, a zoologist whose scrupulona 
occnracy was known to many persons, told me the following 
story of which he was himself an eye-witness ; at the Capo 
of Good Hope an ofBcer had often plagued a certain baboon, 
and the animal, seeing him approaching one Sunday for 
parade, poured water into a hole and hastily made some thick 
rowd, which he skilfully dashed over tho officer as he passed 
by, to the amusement of mony bystanders. For long after- 
wards the baboon rejoiced and triiunphcd whenever he saw his 

■ For the evidrnce on this ' All the rollowiog etatementii, 

kaad, see Mr. J. Traherne Mog- given on the authority of these two 
gridge'a most inlorestiais: wnrk. ii;ituraliatB,Bre Isken fram Rengger's 
■ HarTestlng Ants and Trop-docr 'Naturgcsch. der Siugethie™ voo 
Vle".'1873,p. 12B, 128. Panif nay,' 1830, s. 41-57, aud (rt>a 

' ' Reoharches sur loi Mojnr! dos Brehm's 'Thierlehfn,' B. i 1. 10--87 
Voannis/ 1810, p. 173. 


70 . riu. Dei Hit of M-11 Ia 

Tho iove o[ a dc^ for hi<i master i notorous ns an clJ 
writer quaintly says ° A dog is the oiily tiling on this earth 
" that lUYS you more thin he lu^a himself 

In the agony of death a dog his been known to taross 
his master, and every one has heard of the dog sTifferint' 
under Tivisection, who 1 eked the hand ot tho operator this 
man, unless the operation was fully juitificd by aa imrcTfce 
of our knowledge, or imless he hail a heart of etone mnst haye 
ffilt romorse to the last hoar of his lift 

Ab Whewell '" has ■well asked « ho that rtxds the touching 
" instaneoB of maternal affection related so often of the women ot 
" all nations, and of the females of all animals, can douht that the 
" principle of action is the same in the two cases ? " We see mater- 
nal aBection exhibited in the most trifling details ; thus Rengger 
obserred an American monkey (a Cebus) carefully driving away 
the files which plagued her infant; and Duvaucel saw a 
llylobdies washing the faces of her young ones in a stream. So 
intense is the grief of female monkeys for the loss of their 
young, that it invariably caused the death of certain kinds kept 
under conSnement by Brehm in N. Africa. Orphan monkejs 
were always adopted and carefully guarded by the other monkejs, 
both males and females. One female baboon had so capacious 
a heart that she not only adopted young monkeys of other 
species, but stole young dogs and cats, whieh she eontjmially 
carried about. Her kindness, however, did not go so far aa t« 
share her food with her adopted oSspring, at which Brehm was 
surprised, as his monkeys always divided everything quita 
fairly with their own young ones. An adopted kitten scratched 
this affectionate baboon, who certainly had a fine intellect, for 
she was much astonished at being scratched, and immediately 
examined the kitten's feet, and without more ado bit off the 
claws." In the Zoological Gardens, I heard from the keeper 
that an old baboon ((?. c'racnw) had adopted a Ehesua monkey; 
but when a young drill and mandrill were placed in the cage, 
she seemed to perceive that these monkeys, though distinct 
species, were her nearer relatives, for she at once rejected the 
Ehesus and adopted both of them. The young Ehesus, as I saw, 
was greatly discontented at being thus rejected, and it would, 
like aimughtycbild,annoyandattack the young drill and mandrill 

ited bj Dr. Lander Lindsay, 

72), disputes the possibi 

lity of thi 

Physiology of Mind in the 

act as described by Brel 

im, lor th 

Lnimals;" Journal of Mental 

.:,lte of discrediting 

my wori 

,' April 1871, p. 33. 

Therefore I tried, and fo 

und that 

aidgewater Treatise,' p. 263. 

could readily eeiie wit 

orinv, without any grounds 

teeth the sharp little i 

iinm of 

erlj Ilcview.' July 38J1, p. 

kitten neavly five weeks 



Chaf, in. Mental Powers. 71 

wheneyer it could do so with safety ; this conduct esciting groat 
indignation in the old baboon. Monlejs will also, accordicg to 
Erehm, defend their master when atlaeted by anyone, as well as 
dogs to whom they are attached, from the attacks of other dogs. 
But we here trench on tiie subjects of sympathy and fidelity, to 
which I shall recur. Some of Crehm's monkeys took much 
delight in teasing a certain old dog whom they disliked, as 
well as other animals, in various ingenious waju 

Most of Ihe more complex emotions are common to the 
higher animals and onrselves. Every one has seen how jealous 
a dog is of his master's affeotjon, if lavished on any other 
creature ; and I have observed the same fact with monkeys. 
This shews that animals not only love, but have desire to be 
loved. Animals manifestly feel emulation. They love appro- 
bation or praise; and a dog carrying a basket for his master 
eihibits in a high degree self-complacency or pride. There can, 
I think, be no doubt that a dog feels shame, as distinct from 
fear, and something very like modesty when begging too often 
'.ys food. A great dog scorns the snarling of a little dog, and 
this may be called magnanimity. Several observers have stated 
that monkeys certainly dislike being laughed at; end they 
sometimes invent imaginary offences. In the Zoological Gardens 
I saw a baboon who always got into a fimous rage when his 
keeper took out a letter or book and read it aloud to him ; and 
his rage was so violent that, as I witnessed on one occasion, ho 
bit his own leg till the biood fiowed. Dogs show what may be 
fairly called a sense of humour, as distinct from mere play; if 
a bit of stick or other such object be thrown t« one, he will often 
carry it away for a short distance; and then squatting down 
with it on the ground close before him, will wait until his 
master comes quite close to take it away. The dog will then 
seize it andmsh away in triumph, repeatii^ the same manceuvro, 
and evidently enjoying the practical joke. 

We will now turn to the more intellectual emotions and 
faculties, which are very important, as forming the basis for the 
development of the higher mental powers. Animals manifestly 
enjoy excitement, and suffer from ennui, as may be seen with 
' dogs, and, according to Eengger, with monkeys. All animals 
feel WondeT, and many exhibit Curiosity. They sometimes 
suffer from this latter quality, es when the hunter plays antics 
and thus attracts them ; I have witnessed this with deer, and so 
it is with the wary chamois, and with some kinds of wild-ducks. 
Brehro gives a curious accoimt of the instinctive dread, which 
his monkeys eshibited, for snakes; but their curiosity was 
CO great that they could not desist from nccasionally satiatii^ 


72 The Descent of Man. Pj.bt1 

their horror in a most human BjsMon, by lifting up the fid of tiie 
bos in which, the sniikes ivei'o kept I was so much surprised at 
his accotuit, that I took a stuffed and coi!ed-up snake into the 
montej-houise at the Zoologies] Gardens, and the excitement 
thus caused was one of the most curious spectacles which I eyer 
beheld. Three species of Cercopitheous were the most alarmed ; 
they dashed about their cages, and uttered sharp signal cries of 
danger, which, were Trnderstood by the other monkeys, A few 
young monkeys and one old Anubis baboon alone took no notice 
of the snaka I then placed the stuffed specimen on tho ground 
in one of the larger compartmcnta. After a timeall the monkeys 
collected round it in a lai^ circle, and staring intently, 
presented a most ludicrous appearance. They became extremely 
nervous; so that when a wooden ball, with which they wore 
familiar as a plaything, was accidentally moved in the straw, 
under which it was partly hidden, they all instantly started 
away. These monkeys behaved very differently when a dead 
fish, a mouse," a living turtle, and other new objects were placed 
in their cages ; for though at first fr^htened, they soon 
approached, handled and examined them. I then placed a live 
snake in a paper bag, with the mouth loosely closed, in one of 
the larger compartments. One of the monkeys immediately 
approached, cautiously opened the bag a httle, peeped in, and 
instantly dashed away. Then I witnessed what Brehm has 
described, for monkey after monkey, with head raised high and 
turned on one side, could not resist taking a momentary peep 
into the upright bag, at the dreaifnl object lying quietly at the 
bottom. It would almost appear as if monkeys had some 
notion of zoological affinities, for those kept by Brehm exhibited 
a strange, though mistaken, instinctive dread of innocent lizards 
and frogs. An orang, also, has been kno(vn to be much alarmed 
at the first s^ht of a turtle." 

The principle of Irrdiaivm is strong in man, and especially, as 
I have myself observed, with savages. In certain morbid states 
of the brain this tendency is exaggerated to an extraordinary 
degree ; some hemiplegic patients and others, at the commence- 
ment of inflammatory softening of the brain, unconsciously 
imitate every word which is uttered, whether in their own or in 
a foreign language, and every gesture or action which is per- 
formed near them." Desor" has remarked that no animal 

" I hai 

ivQ given fl short account 

of Mammalia,' J84!, p. 405. 

" Dr. Bateninn ' On Aphm 

ipre^iion of the Emotion^' 

1870, p. 110. 

" Quoted brVogt.'Me'moire 

C L. M.irtin. 'Nat. Hist. 


CBiP. in. ItTenial Powers. 73 

Toluntarily imitates an action perToimcil by man, until in the 
msceuding scale we come to monkeys, wtjcli are well known to 
lie ridiculous mockers. Animals, howsTer, sometimes imitate 
each, other's actions : thus two species of wolves, which had been 
reared by dc^s, learned to bark, as does sometimes the jackal," 
but whether this can be called Tolnnlary imitation is another 
question. Birds imitate the songs of their parents, and some- 
times of other birds; and parrots aro notorious imitators of any 
BOoad which they often hear. Bureau de la Malle gives an 
account " of a dog reared by a cat, who learnt to imitate the 
wsU-known action of a cat licking her paws, and thus washing 
her ears and ftice; this was also witnessed by the celebrated 
naturalist Audouin, I have received several confirmatory ac- 
counts ; in one of these, a dog had not been suckled by a cot, 
but had been brought up with one, togetter with kittens, and 
had thus acquired the above habit, which he ever afterwards 
practised during his life of thirteen years. Bureau de la Malle's 
dog likewise learnt from the kittens to play with a t>all by roll- 
ing it about with his fore paws, and springing on it, A corre- 
spondent assures mo that a cat in his house used to put her paws 
into jugs of milk having too narrow a mouth for her head. A 
kitten of this eat soon learned the same trick, and practised it 
ever afterwards, whenever there was an opportunity. 

The parents of many animals, trusting to the principle of 
imitation in their young, and more especially to their instinctive 
or inherited tendencies, may be said to educate them. We see 
this when a cat brings a live moose to her kittens ; and Bureau 
de la Malle has given a curious account (in the paper above 
quoted) of his observations on hawks which taught their young 
dcsterity, as well as judgment of distances, by iirst dropping 
through the air dead mice and sparrows, which the young 
generally failed to catoh, and then bringing them live birds 
and letttGg them loose. 

Hardly any faculty is more important for the intellectual 
progress of man than Attention. Animals clearly manifest this 
power, as when a cat watches by a hole and prepares to spring 
on its prey. Wild animals sometimes become so absorbed when 
thus engaged, that they may be easily approached, Mr. Bajtifett 
has given me a curions proof how variable this faculty is in 
monkeys. A man. who trains monkeys to act in plays, used to 
purchase common kinds from the Zoolt^cal Society at the pric« 
of five pounds for each ; but he offered to give double the price, 

" 'The Variation of Animals sul " 'Annilea d-s Si-. Snt.' <Ul 

Plants undrr Domesticatkn,' vol. i. Soriw), turn, iiii p. 397. 


74 "^lie Descent of Man Tabt I. 

if lia miglit keep ttree or four of them for a few days, in order 
to select one. WLen asked how he could possibly leani so soon, 
whetter a particular monkey would turn out a good actor, he 
answered that it all depended on their power of attention. If, 
when he was talking and explaining anything to a monkey, its 
attention was easily distracted, as by a fly on the wall or other 
ti'ifling ohject, the case was hopeless. If he tried by pimishment 
to make an inattentive monkey act, it turned eulky. On the 
other hand, a monkey which carefully attended to him could 
always be trained. 

It is almost superfluous to state that animals have excellent 
Memories for persona and places. A baboon at the Capo of Good 
Hope, as I have been informed by Sir Andrew Smith, rocognJecd 
him with joy after an absence of nine months. 1 had a dog who 
\va& savage and averse to ail strangers, and 1 purposely tried his 
memory after an absence of five years and two days. I went 
Dear the stable where he lived, and shouted to him in my old 
manner; ho shewed no joy, hut instantly followed me out walk- 
ing, and obeyed me, exactly as if I had parted with him only 
lialf an bour before. A trainof old associations, dormant during 
live years, had thus been instantaneously awakened in his mind. 
Even ants, as P. Huber" has clearly shewn, recognised their 
fellow-ants belonging to the same community after a separation 
of four months. Animals can certainly by some means judge of 
the intervals of time between recurrent events. 

The Imagination is one of the highest prerogatives of man. 
By this faculty he unites former images and ideas, independently . 
of the will, and thus creates brilliant and novel I'esults. A poet, 
as Jean Paul Eicbter remarks," " who must reflect whether h6 
"shall make a character say yes or no— to the devil with him; 
" he is only a stupid corpse." Dreamii^ gives us the best notion 
of this power ; as Jean Paul again says, " The dream is an in- 
" voluntary art of poetry." The value of the products of our 
imagination deiien»is of course on the number, accuracy, and 
clearness of our impressions, on our judgment and taste in 
selecting or rejecting tho involuntary combinations, and to a 
certain extent on our power of voluntarily combining them. As 
dogs, cats, horses, and probably all the higher animals, even 
birds™ have vivid dreams, and this is shewn by their movements 
and the sounds uttered, we must admit that they possess some 

" ' Les Mteurs des f ouitniB,' 

™ Dr. JerdoQ, ' Birds of 1 

1810, p, 150. t 

ol. i. 1862, p, Hi. Houjeai. 

" QuoledLnDr.Manasley'a'Phy. t 

Fiology and Pithologyof Mind," 18i)3, d 

reamt; 'Faculty Meotiilesi' 


feip. ni. Mental Powers. 7$ 

ftowet of iroagination. There must bo somotliing special, wliich 
tausesdogstohowlintheniglit, and especially duringmooDlight, 
jn that remarkable and melancholj manner called baying. 
All di^s do not do so ; and, according to HouKeau," they do not 
then look at the moon, but at some fised point near the horizon. 
Honzean thinks that their imaginations are disturbed by the 
TBgTie ontlinea of the Eurroundicg objects, and conjiire up before 
tliem fantastic images: if this be so, their feelings may almost 
be called Buperstitlous. 

Of all the faculties of the human mind, it will, I presume, be 
admitted that Reanon stands at the stmimit. Only a few persons 
now dispnte that animals possess some power of reasoning. 
Animals may constantly be seen to pause, deliberate, and resolve. 
It is a Bignificant fact, that the more the habits of any particular 
animal are studied by a naturalist, the more he attributes to 
reason and the less to unleaint instincts.'^ In future chapters 
we shall see that some animals extremely low in the scale appar- 
ently display a certain amount of reason. No doubt it is often 
difScult to distinguish between the power of reason and that of 
instinct. For instance, Br. Hayes, in his work on 'The Open 
Polar Sea,' repeatedly remarks that his dogs, instead of continu- 
ing to draw the sledges in a compact body, diverged and separ- 
ated when they came to thin ice, so that their weight might be 
more evenly distributed. Thia was often the first warning 
which the traTellers received that the ice was becoming thin and 
dangerous. Now, did the dogs act thus from the experience of 
each indiyidual, or from the example of the older and wiser dogs, 
or from an inherited habit, that is from instinct? This instincti 
may possibly have arisen since the time, long ago, when dogs 
were first employed by the natives in drawing their sledges; or 
the Arctic wolves, the parent-stock of the Esquimau! dog, may 
have acquired an instinct, impelhng tliem not t^ attack their 
prey in a dose pack, when on thin ice. 

We can only Judge by the circumstances imder which actions 
are performed, whether they are due to instinct, or to reason, or 
to the mere association of ideas: this latter principle, however, 
is intimateiy connected with reason. A curious case has been 
given by Prof. MSbius,'' of a pike, separated by a plate of glass 
from an adjoining aquarium stocjced with fish, and who often 
dashed himself with such violence against the glass in trying to 

" 'Facnlt^ Meutales des Aci- 1 canDot help thinking, Luwever, 

tnaui,' 1B72, torn. ii. p. IBl. that he goes too far in underrating 

"' Mr. L. H. Morgan's work on the power of Instinct. 
'The American Beaver," 1868, offers " 'Die Bewogooaen dct Thiers 

■ gooif llinstration of this remiik. &c., IBT.S, p. 11. 


76 The Descent of Matt. Pabt I, 

eateh the other fishes, that he was sometimes completely 
Btuanod. The pike went on tima for three mouths; but at last 
learnt contion, and ceased t« do so. The plate of glass was then 
removed, bat the pike would not attack the^e particiilar fishes, 
though he would devour others whicli were afterwards intro- 
duced ; BO strongly waa the i Jta of a Tiolent shock associated 
in hia feeble mind with the attempt on his former neighbours. 
If a savage, who had ne^er seen a largo plate-glass window, 
were to dash himself even once af;amst it he would for a long 
time afterwards associate a shock with a window-frame; but 
very differently from the pike, he nould protiably reflect on the 
nature of the impediment, and be cautious under anali^oua 
circumstances. Now with monkojs, lis we shall presently see, a 
pajnful or merely a disagreeable impression, from aa action onco 
performed, is sometimes sufficient to prevent the animal from 
repeating it. If we attribute this difference between the monkey 
and the pike solely to the association of ideas being bo mueJi 
stronger and more persistent in the one than the other, though 
the pike often received much ite more severe injury, can we 
maintain in the case of man that a similar difference implies the 
possession of a fundamentally different mind ? 

Houaean relates" that, whilst nrossing a wide and arid plain 
in Texas, his two dogs suffered greatly from thirst, and that 
between thirty and forty times they rushed down the hollows 
to search for water. These hollows were not valleys, and there 
were no trees in them, or any other difference in the vegetation, 
and aa they were absolutely dry there could ha be n n 
smell of damp earth. The dogs behaved aa if th y kn w tha 
a dip in the ground offered them the best cban f fi di g 
welter, and Honzeau has often w'tnessed the sam b ha ir n 
other animals. 

I have Been, as I daresay have others, that wl n a m 11 
object is thrown on the ground beyond the reach f n f th 
elephants in the Zoological Gardens, he blows through his trunk 
on the ground beyond, the object, so that the current reflected 
on all sides may drive the object within his reach. Again a well- 
known ethnologist, Mr. Westropp, informs roe that he observed in 
Vienna a bear deliberately making with his paw a current in 
some water, which waa close to the bars of hia cage, so aa to 
draw a piece of floating bread within his reach. These actions of 
(ho elephant and bear can hardly be attributed to instinct or 
inherited habit, as they would be of little use to an animal in a 
state of nature. Now, what is the difference between such 

" 'FacQlt^a Meiit:i1eB dcs AQininoi,' 1873, toin. ii. p. 295. 


Obaf. I!I. Mental Powers. 77 

actions, wten performed by an uncultiTated raaa, and by ono <A 
tile higher aniiaals ? 

The Bavage and the Aog have often fonnd water at a low level, 
and the coincidence under such circmastonces has become asBo- 
ciated in their minds, A cultiTated man would porhax>s make 
some general propoEdtiou on the sutgect ; but from all that we 
know of savages it is extremely doubtful whether they would do 
eo, and a dog certainly would not. But a sarage, as well as a 
dog, would search in the same way, though frequently dis- 
appointed ; and iu both it seems to be equally an act of reason, 
whether or not any general proposition on the subject is 
consciously placed before the mind.'" The same would apply to 
the elephaut and the bear making currents in the air or water. 
The savage ivould certainly neither know nor care by what law 
the desired movements were effected; yet his act would bo 
guided by a rude process of reasoning, as surely as would a 
philosopher in his longest chain of deductions. There would no 
doubt be this difference between him and one of the higher 
animals, that he would take notice of much slighter circum- 
stances and conditions, and would olKcrve any connection 
between them after much less experience, and this would be of 
paramount importance. I kept a daily record of tlie actions of 
one of my infants, aud when he was about eleven months old, 
and before he could speak a single word, I was continually 
struck with the greater quickness, with which all sorts of objects 
and sounds were associated together in his mind, compai-ed with 
that of the most intelligent dogs I ever knew. But the higher 
animals differ in exactly the same way in this power of associa- 
tion from those low in the scale, such as the pike, as well as in 
that of drawing inferences and of observation. 

The promptings of reason, after very short esporience, are well 
shown by the following actions of American monkeys, which 
stand low in thoir order. Iten^er, a most careful obsorvei; 
Slates that when he first gate t^gs to his monkeys in Pai-aguay, 
they smashed them, and thus lost much of their contrails ; after- 
wards they gently hit one end against some hard body, and 
picked off the bits of shell with their fingers. After cutting 
themselves only once with any sharp tool, they would not tonch 
it again, or would handle it with the greatest caution. Lumps 
of sugar were often given them wrapi>ed up in paper; aod 

Prat Huiley has analysed with 

See his BrtiQle, 'Mr. Darwin'a 

irabl 8 clearness the mentaUlaps 

Clitics,' in Ihs 'ConWinporarj Re- 

'hich a man, us well as a dog, 

view,' Nov. 1871, p. 462, nnd in hii 

■Critiq'JH and Esaaja,' 1873, p. 279. 

jgctis to tliat given in my tsit 


?8 Tke Descent jf Man. 

T sometimes put a live wasp in the paptr, so that in 
hastily unfoldiDg it they got stung; after this had once happened, 
they always first held the packet to their ears to detect any 
movement within." 

The following cases relate to dogs. Mr. Colqnhoun" wiugctl 
two wild-ducks, which fell on the further side of 6 stream; hia 
retriever tried to bring over both at once, but could aot succeed; 
she then, though never before known to ruffle a feather, 
deliberately killed one, brought over the other, and returned 
for the dead bird. Col. Hutchinson relates that two partridges 
ii'cre shot at once, odo being killed, the other wounded ; the 
latter ran away, and wns caught by the retriever, who on her 
return came aci'oss tho dead bird; "she stopped, evidently 
" greaily puzzled, and after one or two trials, finding she couid 
" not take it up without permitticg the escape of the winged 
" bird, she considered a moment, then deliberately murdered it 
" by giving it a severe crunch, and afterwards brought away 
" both together. This was the only known instaaco of her 
" ever having wilfully injured any game." Here we have reason 
though not qiiite perfect, for tie retriever might have brought 
the wounded bu;d first and then returned for the dead one, as in 
the case of the two wild-ducks. I give the above cases, as 
resting on the evidence of two independent witnesses, and 
because in both instances tho retrievers, after deliberation, 
broke through a habit which is inherited by them (ttiat of not 
killing the game retrieved), and because they shew how strong 
their reasoning faculty must have been lo overcome a fixed 

I will conclude by quoting a remark by the illustrious 
Humboldt.'' "The muleteers in S. America say, 'I will not give 
" ' yon. the mule whose step is easiest, bnt la, mas racional, — the 
"'one that reasons best;'" and as he adds, "this popula. isprcs- 
" sion, dictated by long eiperience, combats the system of 
" animated machines, better perhaps than all the arguments of 
" speculative philosophy." Nevertheless some writers even yet 
deny that the higher animals possess a trace of reason; and they 
endeavour to esplain away, by what appears to be mere 
"all such fact.; as those above given. 

1 Ni- 

» Mr. Belt, in his 

ing work, 'The KMi 

cai-ngua,' 1874 (j.. 

ii«:>gi'il«sVari0D3 actio 

Cebu^. wbich, 1 thiok 

roBicnirij powe 

'The Hoot ind the I-och,' 

Col. HutcblDson 

ou 'Dog 

?,' 185C 

1, p. 46. 

1 Harratl> 

-',' Eiig. 

,, vol. ii 

i. p. 106.<«l 

to find thi 

net aa 

Mr. Leslie 


ad Divinity, Ek»y> 


'?: 1873, , 

p. 80), in 

Hosted by 


Ch4p. lit. Mental Powers. 79 

It Iws, I think, now been stewn that man and the higliiir \ 
amioals, especially the Primates, have some few instincts in \ 
common. All have the same senses, intuitions, and sensations, — 1 
Kimilar passions, affections, and emotions, even the more comple:£ I 
ones, such as jealoosy, Buspicion, emulation, gratitude, ajid f 
magnanimity ; they practise deceit and are revengeful ; they ai-o I 
sometimes susceptible to ridicule, and even have a sense of / 
humour ; they feel wonder and curiosity ; they possess the same [ 
faculties of imitation, attention, deliberation, choice, memory, \ 
imagination, the association of ideas, and reason, though in very , 
different degrees. The individuals of the same species graduate 
in intellect from absolute imbecility to high excellence. They 
are also liable to insanity, though far less often than in the cnso 
of man."' Nevertheless, many authors have insisted that man is 
divided by an insuperable barrier from all the lower animals in 
bis mental faculties. I formerly made a collection of abovo a 
score of sach aphorisms, but they are almost worthless, as their 
wide difference and number prove the difficulty, if not the im- 
possibility, of the attempt. It has been asserted that man alone 
is capable of progressive improvement ; that he alone makes use 
of tools or fire, domesticates other animals, or possesses property ; 
that no wnimil has the power of abstraction, or of forming 
general concepts, is self-conscious and comprehends itself; that 
no animal employs language; that man alone has a sense of 
beauty, is liable to caprice, tiaa the feeling of gratitude, mystery, 
&c.; believes in God, or is endowed with a consdi 
hazard a few remarks on the more important and ii 
these points. 

Archbishop Sumner formerly maintained" that man alone is 
capable of progressive improvement. That he is capable of 
incomparably greater and more rapid improvement than is any 
other animal, admits of no dispute; and this is mainly 
due to his power of speaking and handing down his acquired 
knowledge. With animals, looking first to the individual, every 
one who has had any experience in setting traps, knows that 


g of the scpposod impnasable 

"tui'es. It is dilT-oult to ul 

between the minds of man 

"stand how anyl>ody who has 

i lower animals, says, "The 

" kept a dog, Of seen an elep 

LCtions, indeed, which have 

"can have any donbti as t 

" animal's power of performin. 

no better foanJatioa than a 

" essential processes of reasonin 

many other metaphysical 

"See -Madness tn Animals 

Dr. W. Under Lindsay, in'Joi 
uf Mental Science,' July 1871. 

things diffei'eat names, they 

■■ Quoted bySirCLyell,'. 

qnityofMan,'p, 497. 


The Descent of Man. I'iST 1 

jouDg anid&ls can bo caught much more easilj than old oucs; 
a,nd they can bo laucli more easily approached by aa enemy. 
Even with respect to old animala, itis impossible to catcli many in 
Lhe same place and in the same kind of trap, or to destroy them 
by the same kind of poison ; jet it is improbable that all should 
jutvo partaken of the poison, and impossihio that all should have 
heou caught in a ti'ap. They must learn caution by seeing their 
hretiirca caught or poisoned. In North America, where the fur- 
boaring animals have long been pursued, they exhibit, according 
to the unanimous testimony of all obserrers, an almost incrediUlu 
amount of sagacity, caution and cunning ; bat trapping has been 
there BO Song carried on, that inheritance may po&eibly have come 
into play. I have reeoiYcd several accounts that when telegrajihs 
a™ first set up in any district, many birds kill themselves by 
Bying against the wires, but that ia the course of a very few 
years they learn to avoid this danger, by seeing, as it would 
appear, their comrades killed.^' 

If we look to saecessive generations, or to the race, there is no 
doubt that birds and other animals gradually both acquire and 
lose caution in relation to man or other enemies f^ and this 
caution is certainly in chief part an inherited habit or instinct, 
but in part the result of individual experience. A good observer, 
Loroy," states, that in districts where foxes are much hunted, 
the young, on first leaving their burrows, are incontestalily much 
more wary than the old ones in districts where they are not much 

Our domestic dogs are deseended from wolves and jackals,'' 
and though they may not have gained in cunning, and may have 
lost in warineBS and suspicion, yet they have progressed in 
certain moral qualities, such as in affection, trust-worthiness, 
tamper, and prolKibly in general intelligenca The common rat 
has conquered and beaten several other species throughout 
Europe, in parts of North America, New Zealand, and recently in 
Pormosa, as well as on the mainland of China. Mr. Swinhoe," 
who describes these two latter cases, attributes the victory of tho 
common rat over the large Nvs aminga to its superior cunning; 
and this latter quality may probably bo attributed to the habitual 

" For additional evidence, with 

» 'r.ettresPhil.siirrJnMllL 

«et,iila, see M. Hoozeau, ' I*> 

des Auiniam,' uouvelle edit. 

Caeultdi Mentales,' torn. ii. 1872, 

p. 88. 

p. U7. 

" See the evideooe on this 

" See, with respect to binis on 

in chap. i. vul i. ' On the Varl 

oceflBlc islanas, my 'Journal of 

of Aniinala^aud I'Lints undo] 

Researches during the rovage of the 

"Beagle," '1845, p, 398, 'Origin 

■* 'Proc. Zoclcg. Soc.' IS-] 

of Spe>:iei,' Stb edit. p. '260. 



Mental Powers. 

exercise of all its fectUtiea in avoiding extirpation by man, ns 
well as to nearly all the less cunning or weak-miiided rats having 
teen continuously destroyed by him. It is, however, possible 
that the success of the common rat may be due to its having 
possessed greater cunning than its feUow-species, before it 
became associated with man. To maintain, independently of any 
direct evidence, that no animal during the course of j^es has 
progressed in intellect or other mental fecultiea, is to beg the 
question of the evolution of species. "We have seen that, ac- 
cordiDg to Lartet, eitisting mammals belonging to several oniers 
have larger brains than their ancient tertiary prototypes. 

It has often been, said that no animal uses any tool; but 
the chijnpanzee in a state of nature craeks a native fruit, some- 
tvhat like a walnut, with a stone.*' Eenggor™ easily taught an 
Amerieau monkey tiins to break open hard palm-nuts ; and 
afiecwards of its own accord, it used stones to open other kinds 
of nuis, as well as boxes. It thus also removed the soft lind of 
fruit that had a dist^eeable flavour. Another monkey waa\ 
taught to open the lid of a large bos with a stick, and after- 
, wards it used tho stick as a lever to move heavy bodies ; and ! 
have myself seen a young orang put a stick into a crevice, slip 
his hand to tne other end, and use it in the proper manner as a 
lever. Tho tamed elephants in India are well known to break 
off branches ol trees and use them to drive away the flies ; and 
this same act has been observed in an elephant in a state ol 
nature." I have seen a young orang, when she thoi:^ht she was 
going to be whipped, cover and protect herself with a blanket or 
straw. In these several cases stones and sticks were employed 
as implements ; but they are likewise used as weapons. Brehm" 
states, on the authority of tba well-known traveller Schimper, 
that in Abyssinia when the baboons belongiag to one species 
((7. jfe^wJa) descend in troops from the moun fains to plunder the 
fields, they sometimes encounter troops of another species 
(C. hawiidryas), and then a tight ensues. The Geladas roll 
down pwit stones, which the Hamadryas try to avoid, and then 
both species, making a great uproar, rush furiously against each 
other. Erehm, when, accompanying the Duke ol Coburg-Gotha, 
aided in an attack with fire-arms on a troop of baboons in the 
pass of Monsa in Abyssinia. The baboons in return rolled so 
many stones down the mountain, some as large as a man's head, 
that the attackers had to beat a hasty retreat ; and the pass wiw 

" The ' Indian FieU,' March *, 
• 'Tliierloben,' B. i, ». 73, 82. 


S2 The Descent of Man. Pakt I, 

(iplualiy closed for a time against the caravan. It dcBerren 
notice that these baboons thus acted in concert. Mr. 'Wallaco" 
on tbree occasioas saw female orangs, accompanied by their 
young, " breaking off branclies and the great spiny froit of the 
■' Durian tree, with every appearance of rage; causing Buch a 
"shower of missiles as effectually kept us from approaching too 
" near the tree." As I have repeatedly seen, a chimpanzee will 
throw any object at hand at a person who offends him; and the 
before mentioned baboon at the Cape of Good Hope prepared 
mud for the purpose. 

In the Zoological Gardcng, a monkey, which had weak teeth, 
used to break oiren nuts with a stone ; and I was assured by the 
keepers that after using the stone, he hid it in the straw, and 
would not let any other monkey touch it. Here, then, we have 
the idea of property ; but this idea is common to every dog with 
a bone, and to most or all birds with their nests. 

The Duke of Argyll" remarks, that the fashioning of an 
implement for a special purpose is absolutely peculiar to man ; 
and he considers that this forms an immeasurable gulf between 
him and the brutes. This is no doubt a very important dis- 
tinction ; but there appears to me much truth in Sir J. Lubbock's 
suggestion,** that when primeval man first used flint-stones for 
any purpose, he would have accidentally splintered them, and 
would then have used the sharp fragments. From this step it 
would be a small oue to break the flints on piirpose, and not a 
very wide step to fashion them rudely. This latter advance, 
however, may have taken long ages, if we may judge by the 
immense interval of time which elapsed before the men of the 
neolithic period took to grinding and polishing their stone tools. 
In breaking the flints, as Sir J. Lubtwck likewise remarks, 
sparks would have been emitted, and in grinding them heat 
-*ould have been evolved : thus the two usual methods of 
" obtaining fire may have originated." The nature of fire would 
have been known in the many volcanic regions where lava 
occasionally flows through forests. The antjiropomorphoue 
apes, guided probably by instinct, build for themselves tcra- 
poraty platforms ; bnt as many instmcts are largely controlled 
by reason, the simpler ones, such as this of building a platform, 
might readily pass into a voluntary and conscious act. The 
orang is known to cover itself at night with the leaves of the 
Pandanus; and Brehm states that one of his baboons used to 
protect itself from the heat of the sun by throwing a straw-mat 

*i 'The Malay ArohipcUgo,' vol. 145, 147. 
i. 1869, p. 87. " ■ Prehistoric Times,' 1865, p, 

*• ' Primeval Man,' 18fi9, pp. 473, &c 


Oa-'T-. in. Menial Powers. S3 

ovpr ite head. In these eeTeral huliits, we probably eeo tiie first 
stops towards some of the simpler arts, such as tade aichitectrire 
fluii dress, as thej arose amongst the early pti^mtors of man. 

^Ssirocffon, Otn&nd Coac^tiom, tielf-cnnsciotisness. Menial 
Individuality. — It would be very diffleult for anj one with even 
much more knowledge than I possess, t« determine how fer 
animals exhibit any traces of these high mental powers. This 
difaoulty arises from the impossibihty of judging what passes 
through the mind of an animal ; and again, the fact that writers 
differ to a great extent in the meaning which they attribute to 
the above terms, causes a further difBciilty. If one may judge 
from various articles which have been published lately, the 
greatest stress seems to be laid on the supposed entire absence 
in animals of the power of abstraction, or of forming general 
concepts. But when a dog sees another dc^ at a distance, it is 
often clear that ho perceives that it is a dog in the abstract ; for 
when he gets nearer his whole mamier suddenly changes, if the 
other d(^ be s. friend. A recent writer remarks, that in all suc'i 
cases it is a pure assumption to assert that the mental act is 
Dot essentially of the same nature in the animal as in man. It 
either refers what he perceives with his senses to a mental 
concept, then so do both." When I say to my terrier, in an 
eager voice (and I have made the trial many times), " Hi, hi, 
where is it?" she at once takes it as as%n that something is to 
be hunted, and generally first looks quickly all around, and 
then rushes into the nearest thicket, to scent for any game, but 
finding nothing, she looks up into any neighbouring tree for a 
squirrel. Now do n t th act n ! arly shew that she had in 
her mind a general la pt that some animal is to be 

discovered and hunted 

It may be freely dm tted that n animal is self-oonscious, 
if by this term it is miplied hat he reflects on such points, be 
whence ha comes or wh th h will g , or what is life and death, 
and BO forth. Bat h w can w feel ure that an old dog with an 
oicellent memory and some power of imagination, as shewn by 
his dreams, never reflects on his past pleasures or pains in the 
chase ? And this would be a form of self-conBciousness. On the 
(rther hand, as Biichner" has remarked, how little can the hard- 
worked wife of a degraded Australian savage, who uses very 
faw abstract w<r rds. and cannot count above four, exert her self- 
COBBcionsness, or reflect on the nature of her own existence. It 
IE generally admitted, that the higher animals possess memory, 

" M(. Hookham, in a letter to " 'Conferences snr la Tliiortt 

Pn>£ Mai Mtlller, in th; ' Birmisg- Datwbienne,* French traualat. 
bam KewB,' May 1S73, JBSB p. 132. 


84 The Dt.Si.uii of Man. Pabt I. 

attentinn, uRSociation, and oven some imi^jtiatjon and reason 
If tteso powers, which differ much m diffL-rent amraah, aro 
capable of improToiaent, there seemi no great impiobabilitj' m 
more complei fiuinlties, bbcJi as the hifther forms of abstraction, 
anii self-conaciausncBS, &o., having been evolved through tho 
development and combination of the simpler one^ It hns been 
u^ed againRt the views hero maintamed, that it i? impossible 
to say at whitt point in the aswindiug scale animals become 
capable of abstraction, &c. ; but who can say at what age this 
occurs in our yr^ng children ? We see at least that such powers 
ire developed in children by imperceptible degrees 

That animals retain their mental individuality is unqiiostion 
able. 'When my voi^e awakened a train of old associations lu 
the mind of the before-mentioned dog he mu'rt have reta ne ! 
his mental individnality, although every atom ot his bram had 
probably undergone change more than ones during the interval 
of iive years. This dog might have brought forward the 
argument lately advanced to crush all evolutioruhts and said 
" I abide amid all mental moods and all material changes 
"The teaching that atoms leave their rmpre^sions as legacies to 
"other atoms falling into the places they have vacate 1 k con 
" tradiatory of the utterance of consciousnein and is therefore 
" false ; but it is the teaching necessitated by evolutionism, con- 
" sequently the hypothesis is a false one," " 

Langvage.—Tbis tk;Dlty has Justly been considered as one of 
the chief distinctions between man and the lower ajiimals. But 
man, as a highly competent Judge, Archbishop Whately remarks, 
" is not the only animal that can make use of language to express 
" wliat is passing in his mind, and can understand, more or less, 
"what is so espressed by another."" In Paraguay the Cebun 
asaTix when eicited titters at least six distinct sounds, which 
esdto in other monkeys similar emotions." The moTementa of 
the features and gestures of monkeys are understood by us, and 
tbey partly understand ours, as Bengger and others declare. It 
is a more remarkable fact that the dog, since being domesticated, 
has leamt to bark" in at least four or five distinct tones. 
Although barking is a. new art, no doubt the wild parent-species 
of the Aog eipressed their feelings by erica of various kinds. 
With the domesticated dog we have the bark of eagerness, as in 
theehase; thatof anger, asweli as growling; the yelp or howl ot 
despair, as when shut up ; the baying at mght ; the bark of Joy, r-s 

« Tne Rev. Dr. J. M'Cann, ' Anli- •• Kem^Ter, ibid. s. 45. 

Darwinism,' IBSfl, p. 13. " S*9 mr 'Vnriation of All- 

*■ Qontud in 'AnthropologicRl Re- mala Bod Plants under Domwli™. 

rt«w • 16C4, p. 158. ticn," vol. i, p, 27. 


OHAP. lit. 

Mental Powers. 


(Fhen starting on a walk with his mastoi ; and the very diBtinct 
one of demand or supplication, as when wishing for a door or 
window to be opened. According to Houzeau, who paid par- 
ticular attention to the subject, tbo domestic fowl utters at least 
a dozen Eignificant Eonnds.™ 

The habitual use of articulate language is, howerer, peculiar 
to loan ; but be uses, in common with the lower animajs, inarti- 
culate cries to express hia meaaiag, aided by gestures and the 
movements of the muscles of the Ihce.^' This especially holds 
good with the more simple and rivid feelir^, which are but 
little conuectwl with our higher inteliigeuce. Our cries of paiu, 
fear, surprise, anger, together with theii appropriate actions, 
and tlie murmur of a mother to her beioyed child, are raoro 
oxpressiTe than any words. That which distingujslies man 
from the lower animals is not the undetstandii^ of articulate 
sounds, for, as OTery one knows, dc^ uudcrstand many words 
and sentences. In tijs respect they are at the same slage of 
development as infants, lietween the ages of ten and twehe 
months, who understand many words and short sentences, 
cannot yet utter a single word. It is not the mere articulation 
which is our distinguishing character, for parrots and other 
birds possess this power. Nor is it the mere capacity of c 
necting definite sounds with definite ideas; for it is certain that 
some parrots, which have been taught to speak, connect n 
erringly words with things, and persons with events.^ The 
lower animals differ from man solely in his almost infinitely 
larger power of associating together the most diversified I 

" 'Faculty Mentalea des Ani- 
maui,' Mm. ii. 1873, p. 346-349. 

in Mr. E. B. Tylor's very iDtetesting 
work, 'Resaacches into the Early 
History of Mankinil,' IStifi, chaps. 

" 1 have iweived se 

1I detailed 

Sir J. SullTau, whom I know 

«u African parrot, long kepC in 
(athet's house, invariably cal 
certain persons of tlie household, 

daid **good morning" to every one 
bmakfeat, and " good night " lo ei 
as they left the room at night, b 

Tn Sir J. Snlivan's father, hs ui 


t sentence, whiot was never 
repeated after his. father's 
h. He BColded violently a 
]ge dog which came into the 
1 through the open windon-; 
he ecoJded another parrot (say- 
"jou nanghtf polly") which 

,0 persons arriving, and " good- 

>ye, old fellow," to tjioee departing. 

couW add several other such 


S6 The Descent of Man. I'^^'i' ^■ 

I dounde and ideas; and this obTionslj depends an th* high 
development of his mental powers. 

As Home Took, one of the founders of the noble science o£ 
philology, observes, language is an art, lite brewing or babjng ; 
but writing would have been a better simile. It certainly is not 
a true instinct, for every language has to be learnt. It diffaifi, 
however, widely from all ordinary arts, for man has an in- 
stinctive tendency to speak, as we see in the babbie of our 
yoni^ children ; whilst no child has an instinctive tendency lo 
brew, bake, or write. Moreover, no philologist now supposes 
that any language has been deliberately inTented; it has been 
slowly and unconsciously developed by many steps." The 
sounds uttered by birds offer in several respects the nearest 
analc^ to language, for eII the members of the same species utter 
the same instinctive cries espressive of their emotions; and all 
the kinds which sing, exert their power instinctively; but the 
actual 601^, and even the call-notes, are learnt from their 
parents or foster-parents. These sounds, aa Daines Barrington" 
has proved, "are uo more innate than language is in man." 
The first attempts to sing " may bo compared to the imperfect 
" endeavour in a child to babble." The young males continua 
practising, or as the bird-catchers say, " recording," for ten or 
eleven months. Their first essays show hardly a rudiment of 
the future song ; but as they grow older we can perceive what 
they are aiming at; and at last they are said "to sing th«ir 
" song round." Nestlings which have learnt the song of a distinct 
species, as with the canary-birds educated in the Tyrol, teach 
and transmit their new song to their offspring. The slight 
natural differences of song in the same species inhabiting 
different districts may be appositely compared, as Barrington 
remarks, "to provincial dialects;" and the soi^ of allied, 
though distinct species may be compared with tbo languages of 
distinct races of man. i have given the foregoing details to 
sbew that an instinctive tendency to acquire an art is not 
peculiar to man. 

With respect to the origin of articulate language, after having 
read on the one side the highly interesting works of Mr. Heus- 

" Ses soma good remarks on this " gard h mmed ud tt be 

head by Prof. Whitnej-, io his " atla n d u asreg^inl' 

■Orient^ and Liugnistic Studion,' "the h nse^ a s of tlie 

1873, p. 35*. He ohsecves that the " iol. 

in the deielcpmeat of la^^un; 
" norks both cunsciouely arid n 
" oHuciinuly ooDScionsly as i 


Chap. IIL Mental Poivers. 8; 

leigh 'Wedg^vood, the Eer. P, rarrftr, aad Prof. Schleichei," aiui 
the celebrated lectures of Prof. Mas MUller ort tte otter side, I 
cannot doubt that lai^unge owes its origin to the imitation and 
modification of various natural sounds, the voices of other 
animals, and man's own instinctive cries, aided by signs and 
gestures. When we treat of sesnal selection we shall see that 
pnmoTal man, or rather some early progenitor of man, probably 
first used his voice in producing true musical cadences, that is 
in singing, as do some of the gibbon-apes at the present day ; 
and we may conclude fiom a widely-spread analogy, that this 
power would have been especially exerted during the courtship 
of the seses,— would have expressed various emotions, such as 
love, jealousy, triumph, — and would have served as a challenge to 
rivals. It is, therefore, probable that the imitation of musical 
cries tj articulate sounds may have given rise to words expres- 
sive of various complex emotions. The strong tendency in our 
nearest allies, the monkeys, in microcephalous idiots," and in 
the barbarous races of mankind, to imitate whatever they hear 
deserves notice, as bearing on the subject of imitation. Since 
monkeys certainly understand much that is said to them by 
man, and when wild, utter pignai-cries of danger to their 
fellows; " and since fowls give distinct warnings for danger ot 
the ground, or in the sky from hawks (both, as well as a third 
cry, intelligible to dt^),"" may not some unusually wise ape-like 
animal have imitat«i the growl of a beast of prey, and thus 
told his fellow-monbejs the nature of the expected danger ? This 
would have been a first step in the formation of a language. 

As the voice was used mote aiid more, the vocal organs would 
have been strengthened and perfected through the principle of 
the inherited effects of nse ; and this would have reacted on the 
power of speech. But the relation between the continued use of 
language and the development of the brain, has no doubt been 
far more important. The menial powers in some eaily pro- 
genitor of man must have been more highly developed than in 

" 'On the Origin of Langoage,' 

» Vost, • M^oire sur lei Micro. 

bj H. WedgwoQd, 1866. 'Chapters 

c^phalea,' 1SH7, p. 169. With re- 

on Language," by the Rei-. F. W. 

spect to siiTages, 1 have given some 

FaiTar, 1865. These works ars 

facts in my ■ Journal of Besearehes,' 

most interealio?. See also ' Do la 

&0., 1846, p. 20S. 

Phjs. et de I'arole,' i>ar Albert 

" See clear evidence en this head 

Lemoine, 1865, p. 190. The worlt 

in the two works so often quoted, 

no this subject, by the Ute Prof. 

by Brehm and Rengger. 

Aog. Schleicher, has been translated 
by Dr, BikherB into English, under 

" Hou.ean gives a very curious 

account of his observations on tilii 

lb* title of • Darwinism tosted by 

lubject in his < f acultt^ Mentalca 

the &asacs of iangu.-ige,' 1869 



the Descent of Mai. 

TAai T. 

any esisting ape, before eTen the most imperfect form of speocli 
eouJd have come into use; but we may confidently believe that 
the contiuued use and advaneemeat of this power would have 
reacted on the mind itself, by enabling asd encouraging it t^i 
carry on long tiadns of thought. A complex train of thought 
can no more be carried on without the aid of words, whether 
Hpokeii or silont, than a long calculation without the use of 
figures or algebra. It appears, also, that ovea an ordinery train 
of thought almost requires, or is greatly facilitated by some 
form of lou^age, for the dumb, deaf, and blind girl, Laura 
Bridgmau, was obseived to use her fingers whilst dreaming.'"' 
/ Nevertheless, a long succession of vivid and connected ideas may 
/ pass tlirough the mind without the aid of any form of language, 
OS we may infer from the movements of dc^ during their 
dreams. We have, also, seen that animals are able to reason 
to a certain estent, manifestly without fJie aid of language. 
The intimate connection between the brain, as it is now 
developed in us, and the faculty of speech, is well shewn by 
those curious cases of brain-disease in which speech fc specialiy 
affected, bk when the power to remember substantives ia lost, 
whilst other words can be correctly used, or where substantives 
of a certain class, or all except the initial letters of substantives 
and proper names are forgotten.'" There is no more improl)- 
ability ia the continued use of the mental and vocal organs 
leading to inherited changes in their structure and functions, 
than in the case of handwiiting, which depends [partly on the 
form of the hand and partly on the disposition of the mind ; and 
hand- writing is certainly inherited." 

Several writers, more especially Prof. Max Miiller,'' have 
lately insisted that the use of language implies the power of 
forming general concepts ; and that as no animals are supposed 
to possess this power, an impossible barrier is formed between 
them and man.'' %Yit)i respect to animals, I have already 

" See remurka ou tl^is head by 

« Lectures on'Mr.Darwin'sPhi- 

l>r. Maudiley, 'The Physiologj and 

loaophy of l^anguage,' 1873. 

I'athDlogj- of Mind,' 2di1 edit. 1868, 

" Tbe judgment of a dislin- 

p. 199. 

gnished pbilologist, such as Vroi. 

" Many cm-iom oases have been 

Whitney, will have for more weiglil 

recorded. See, 6r instance. Dr. 

on tllis point than anything that 

riateman ' On Aphasia,' ISTO, p. 27, 

I can say. He i^emarks (' Oriental 

31, 53, 100. &o. Ako, 'Inquiriei 

and Linguistic Studies," 1873, p 

Concerning the InMlleotual Powers, 

297), in speaking of Bleek's Tiews . 

fcjr Dr. Abeccrombie, 1838, p. 150. 

" "fhe V^iriation of Animai? 

and J'lanls under Domestication,' 

" of tnougbt, indispensable to the 

lol, ii. p. e. 

"development of the p«wer <A 


t3HAP, m. Mental Powers. 8g 

ondeavoured to show that the? have this power, at least in a 
rude and incipient degree. Ab &r as concerns infftnta of from 
ten to eleven months old, aad deaf-mutes, it seems to me in- 
credible, that thej should be able (o connect certain sounds with, 
certain genei'al ideas as quickly as they do, unl&sa such ideas 
were alr«tuiy formed in their minds. The same remark may ho 
extended to the more intelhgent animals ; as Mr. Leslio Stephen 
observes,"' " A dog fraToes a general concept of cats or shoep, 
" and knows the corresponding words as well as a philosopher. 
" And the capacity to nndecatand is as good a proof of vocal 
" intell^ence, though in an inferior degree, as the capacity to 
" speak." 

Why the Cleans now used for speech should have been 
originally perfected for this purpose, rather than any other 
oi^ns, it is not difficult to see. Ants have considerable powers 
of intercoQiTOunication by means of their antennsE, as shewn by 
Hubcr, who devotes a whole chapter to their language. Wo 
might have used our fingers &a efficient instj;uments, for a 
person with practice can report to a deaf man every word of a 
speech rapidJy delivered at a public meeting; but the loss of 
our hands, whilst thus employed, ironld have been a serious 
inconvenience. As all the higher mammals possess vocal organs, 
constructed on the stme generoJ plan as ours, and used as a 
muic5 of oommnmcat on it was obviously probable that these 
same organs would bo Bt 11 further developed if the power of 
commnn cation had to be improved and tliis has been eSccted by 
the a d of adjamng and veil adarted parts, namely the tongue 
and lips"* ihe fact of the higbtr apes cot usmg their vocal 
organs for speech n> doubt dtptJids on their mtelhgence not 
1 iving been fcufficicntly advanced. The possession by them ol 

h Dk Dg to the d 3t nctDt.B and 

finge.^^ into mutation of spoken 

ret. ant gnpUllty ol coga 

woids" Mai Milller giros .a 

ons to tha iul mastery of <. n 

talcs (' on Mr Darwin. 

Ph loaophT of L,iagiJjge,' IdTd, 

ua make thought aboolutelv m 

th rd lecture) the followmg aphor- 

osi ble mtli ut speech idsntlfj- 

m " There is no thought witb- 

ig tha facn ty w Ih ts iBBtru 

ont woids, as little as there ir» 

ent Ha m ght just «. leasoo 

words without thouglit " Wbii 

bly assort that the hnman hand 

innot act without a tool With 

g ea to the word thcnght I 

uch a dootr ua to ita I from he 

" ' lasays on Fiee-th-aking,' &c, 

annot stop ehoit of Uullera 

18-^ p 82 

orst paradoias, that an inf n 

' See some food remarks to tJii> 

It /a « not peak ng^ la i ot a 

efloct. by Dr. Maudsley, 'The Phy- 

am a be o" and that deaf mutes 

-ology ana Vathoiogy of Mind, 

not be ome puiS ssed of eason 

I8b8, p. 199. 

ntu they laara o twist their 


go T/ie Descent of Mem. 

organs, which with long-cotitinued practicB might have been 
used for speech, although not thus used, is paralleled by the 
CQse of manj birds which possesa organs fitted for siting, 
though they never sing. Thus, the nightingale and crow have 
vocal oigans similarly constructed, these being used by tha 
former for diversified song, and by the latter only for croaking." 
If it be asked why apes have not had their intellects developed 
to the ^me degree as that of man, general causes only can be 
■^signed in answer, and it is unreasonable to expect anything 
more definite, considering our ignorance with respect to the 
successive stages of development through which each creature 
has passed. 

Tho formation of different languages and of distinct species, 
and the proofe that both have been developed through a gradual 
process, are curiously parallel." But we can trace the fonnation 
of many words further back than that of species, for we can 
perceive how they actually arose from the imitation of various 
sounds. We find iu distinct languages striking homologies due 
to community of descent, and analogies due to a similar process 
of formation. The manner iu which certain letters or sounds 
change whsu others change is very Uke correlated growth. Wo 
have in both oases the reduplication of parts, tho effects of long- 
eoiitimied use, and so forth. The frequent presence of rudi- 
ments, both in languages and in species, is still more remarkable. 
Tho letter m in the word aw., means i ; so that in the expres- 
sion / am, a superfluous and useless rudiment has been retained. 
In the spelling also of words, letters often remain as the rudi- 
ments of ancient forms of pronunciation. Languages, like 
oi^anio beings, can be classed in groups under groups; and they 
nan be classed either naturally according to descent, or arti- 
ficially by other characters. Dominant languages and dialects 
spread widely, and lead to the gradual extinction of other 
tongues. A language, like a species, when once extinct, never, 
as Sir C. Lyell remarks, reappears. The same language never 
has two birth-places. Distinct languages may be crossed or 
blended tt^ther,"' We see variability in every tongue, and new 

Ma=gLlli.rar, ' Hist, of British 

display any nnusiul capadty for 

1= vol 11 1839 i 29 Ad 

llent obserrei Mr Blackwall, 

1S34, p. 158. 

a\a that tha magp a leirns to 

ounce BiDgle wuids and even 

rallellsm beta-eell tlia development 

t seutenees more rcidilv 

ftt any other Erit si bird jet. 

Sir C Lyell in 'The Qeolog. Evi- 
dences of the Antiquity of M«n,' 

e adda aftei long ino (.losely 

ttigating Its habits he has 

1863, ehap. iiiii. 

" See remarks to this cflect bj 


CBAr. III. Metital Powers. ^l 

words are continually cropping np; but aa there is a limit t*) 
tiie powers of the memory, single words, like whole languages, 
gradually become extinct. As Mas MuUor" has well re- 
marked ;— " A stri^Ie for life is constantly going on amongst 
" tbo words and grammatical forms in each language. The 
" better, the shorter, the easier forms are coustantJy gaining the 
"upper baJid, and thej owe their success to their own inherent 
" virtue." To ttese more important causes of the survival of 
certain words, mere novelty and feshion may be added ; for 
there is in tlie mind of man a stror^ love for slight changes in all 
things. The survival or preservation of certain fevonred words 
in the stm^le for osistenee is natural selection. 

The perfectly i:egular and wonderfully complei construction 
of the languages of many barbajous nations has often been 
advanced as a proof, either of the divine origin of these lan- 
guages, or of the high art and former civilisation of their 
founders. Thus P. von Schlogol writes : " In those languages 
." which appear to be at the lowest grade ot intellectual culture, 
'■ we frequently observe a very high and elaboraifl degree of art 
" in their grarcmatical structure. This is esi)ecially the case 
" with the Basque and the Lapponian, and many of the Ame- 
" rican langnages."^ But it is assuredly an error to speak of 
any language as an art, in the sense of its having been elabor- 
ately and methodieally formed. Philologists now admit that 
conjugations, declensions, &c., originally existed as distinct 
words, since joined together; and as such words express the 
most obvious relations between objects and persons, it is not 
surprising that they should have been used by the men of most 
races during the earliest ages. With respect to perfection, the 
following illustration will best shew how easily we may err: a 
Crinoid sometimes consists of no less than 150,000 pieces of 
shell," all arranged with perfect symmetry in radiating linos ; 
but a naturalist does not consider an animal of this kind as 
more perfect than a bilateral one with comparatively few parts, 
and with none of these pacts alike, excepting on the opposite sides 
of the body. He justly considers the differentiation and special- 
isation of organs as the test of perfection. So with languages ■ 
the most symmetrical and complex ought not to be ranked abovo 
irregular, abbreviated, aaid bastai'dised languages, which have 

the Rev. F. W, Farrar, in an ia- '" Qnot&l by a S. Wake, 'Chap- 

tore^ting article, entitled ' Phili- tei3 od Man,' ISSS, p. 101. 
Iniry aad Darwinism' in 'Nature,' " Buokland, 'Biidgeaater Trea- 

tllrch S4th, 1870, p. 52a tise,' p. 411. 
"s 'Nature,' Jan. Stli,187C p. 257. 


92 The Descent of Man. Pisr L 

borrowed csproBsivo words aod useful fomiB of Ronstmotion from 
■various conquering, conquered, or immigrant races. 

From these few and imperfect remarks I conclude that tho 
extremely complex and regular constmctiou of many barbarous 
languages, is no proof that they owe their origin to a special 
act of creation." Nor, as we have seen, does the faculty lA 
articulato speech in itself offer any insupemblo objection to 
tbe belief that man has been doTcloped from some lower 

Senss of Beauty. — This sense has been declared to be peeuliiir 
to man, I refer here only to the ploasnre given by certain 
colours, forms, and sounds, and which may fiirly be called a 
sense of the beautiful ; with cultivated men such sensations are, 
however, intimately associated with complex ideas and trains of 
thought. When we behold a male bird elaborately displayii^ 
his graceful plumes or splendid colours before the female, 
whilst other birds, not thus decorated, make no such display, 
it is impossible to doubt that she admires the beauty of her 
male partner. As women everywhere deck themselves with 
these plumes, the beauty of such ornaments cannot be disputed. 
As we shall see later, the nests of humming-birds, and the 
playing passages of bowcr-hirds are tastefully ornamented 
with gaily-coloured objects; and this shews that they must 
receive some kind of pleasure from the sight of such things. 
With the great majority of animals, however, the taste for the 
beautiful is confined, as far as wo can juc^, to the attractions 
of the opposite sex. The sweet strains poured forth by many 
male birds during the season of love, are certainly admired by 
the females, of which fact evidence will hereafter be given. If 
female birds had been incapable of appreciating the beautiful 
colours, the ornaments, and voices of their male partners,all the 
labour and anxiety exhibited by the latter in displaying their 
charms before the females would have been thrown away; and 
this it is impossible to admit. Why certain bright colours 
should excite pleasure cannot, I presume, he explained, any 
more than wby certain flavours and scents are agreeable; but 
habit ha£ something to do with the result, for that which is at 
first unpleasant to our senses, ultimately becomes pleasant, and 
habits are inherited. With respect to sounds, Helmholtz has 
explained to a certain extent on physiological principles, why 
tarmonies and certain cadences are agreeable. But beside 
this, sounds frequently reonrrtng at irregular intervals we 


Chap. III. Mental Powers. 93 

higtly disagreeable, as every one will admit who lifls listened at 
Digtt to the irregular flapping of a rope on hoard ship. The 
Bame principle seems to come into play with vision, as the 
eye prefers symmetry or figures with Bome regular recurrenoe. 
Patterns of this kind aio employed by even tie lowest savages 
as ornaments; and they have been developed tlux)ugh sesual 
selection for the adornment of some malo am'mals. Whether we 
can or not give any reason for the pleasure thus derived from 
vision and hearing, yet man and many of the lower animals aro 
alike pleased by the same colours, graceful shatling and forms, 
and the same Bounds. 

The taste for the beautiful, at least as far aa female beauty is 
concerned, is not of a special nature in the human mjad ; for it 
differs widely in the different races of man, and is not quit* the 
same even in the different nations of the same race. Juicing 
from the hideous ornaments, and tha equally hideous music 
admired by most savages, it might be urged that their sssthetic 
faculty was not so highly developed as in certain animals, for 
instance, as in birds. Obviously no animal would ba capable of 
admiring such scenes as the heavens at night, a beautiful land- 
scape, or refined music ; but snch high tastes are acquired 
through culture, and depend on complex associations ; they are 
not enjoyed by barbarians or by uneducated persons. 

Many of the thculties, which have been of inestimable service 
to man for his progressive advancement, such as the powers of 
the imagination, wonder, curiosity, an undefined sense of beaiity, 
a tendency to imitation, and the love of escitement or novelty, 
could hardly fail to lead to capricious changes of customs and 
fashions. 1 have alluded to this point, because a recent writer ^ 
has oddly fixed on Caprice " as one of the most remarkable and 
"typical differences between savages and brutes." But not 
only can we partially understand how it is that man is from 
various conflicting influences rendered capricious, but that 
the lower animals are, as we shall hereafter see, likewise ciipri- 
oious in their affections, aversions, and sense of beauty. There 
is also reason to suspect that they love novelty, for it own saio. 

Bdief in Goii^-Rdigion. — There is no evidence that man was I 
aboriginally endowed with the ennobling belief in the existence 
of an Omnipotent God. On the contrary there is ample ovidenee, 
aerived not from hasty travellers, but &om men who have locg 
resided with savages, that numerous races have existed, and still 
exist, who have no idea of one or more gods, and who have no 

" 'The Sppctnfor,' Dec 4th, 18G9, p. 1130. 



The Descent of Ma>. 

Past 1, 

trords in their languages to ospress euoh an Moa."' The qnostion 
is of course wtoHy diatinct from that higher one, whether there 
eiistK a Creator and Euler of the universe ; and this has b^eu 
answered in theaffirmatiTe by some of the highest intellects that 

, have ever esiBted. 

If, however, we include under the term "religion" the belief 
in unseen or spiritual agencies, the case is wholly different ; for 

; this belief seems to be universal with the less civilised races. 
Nor is it diflScuIt to comprehend how it arose. As soon as the 
important faculties of the imagination, wonder, and curiOHity, 
tc^ther with some power of reasoning, had become parl.ially 
developed, man would naturally crave to understand what was 
passing around him, and would have vagniely speculated on his 
own existence. As Mr. MXennan" lias remarfeed. " Some explan- 
" ation of the phenomena of life, a man must feign for himself- 
" ajid to judge from theunfversalityof it, the simplest hypothesis , 
" and the first to occur to men, seems to have been that natural 
" phenomena, are ascribable to the presence in animals, plants, 
'" and things, and in the forces of nature, of sncli spirits promptiug 

"to a 


is alw probable, as Mr. Tylor has shewn, that dreams may have 
first given rise to the notion of spirits ; for savages do not readily 
distinguish between snhjeciive and objective impressions. When 
a savage dreams, the figures which appear before him are 
believed to have come from a distance, and to stand over him ; 
or " the soul of the dreamer goes out on its travels, and comes 
" Lome with a remembrance of what it has seen."" But imtil 

us of religious belief thionghoiit 
world, by man being led thniueh 
1ms, Ehadows, nnd other c-iases, 

ince, cni'poreil nnd Bpintunl. As 

'. W. 1 

historic Times,' 2nd edit. 1809, p. 
564; and espedally the ehnpters on 
JieligioD in fais 'Ch-igin of Ciyilisa' 

" 'The Worship of Animals and 

He then fmther shews that na 

Plauts,' In the ' Fortnightly Review,' 

o( nicknames given from e- 

Oct. 1, 1869, p. 432. 

animal or othej object, to the « 

•■ Tylor, 'Enrly HUtory of Man- 

progenitors or fonndeis of a tr 

kind,' 1865, p. 6. See also the 

are supposed after a long intoi 

three striking chapters on the Do- 

to ripi'tsent the real progenitoi 

the tribe; and such animal or ob 

' Origin of Qvilisation,' 1870. In a 

is then naiorallj believed still 

like manner Mr. Herbert Spencer, 

eiist as a spirit, is held sacred, 

in his ingenious essay in the ■Fort- 

worshipped as a god. NsTerthe 

nightly Review' (May lat, 1870, 

1 cannot tjut gospect that theii 

p. 6351, accounts for the earliest 

a still earlier and ruder sUge, w 


Chap. Ill, Mental Powers. 9; 

the faculties of iraflginatioii, curiosity, reason, iSe,, tad been 
fairly well deTcloped in the loiiid of man, his dreams would not 
hsYe led Mm to believe in spirits, any more than in tie case of 

The tendency in savages to imagine that natoral ohjects and 
agencies are animated by spiritual or Uring essences, ia perhaps 
illustrated by a little fact which I once noticed : my dog, a full- 
grown aud very sensible animal, was lying on tbe lawn during a 
hot and still day ; bat at a little distance a slight breeze occa- 
sionally moved an open parasol, wliich would have been wholly 
disregarded by the dog, had any one stood near it. As it was, 
every time that the parasol slightly moved, the dog growled 
fiercely and barked. He must, I think, have reasoned to liimself 
in a rapid and unconscious manner, that movement without any 
apparent cause indicated the presence of some strange liviug 
agent, and that no stranger had a right to be on his territory. 

The belief in spiritual agencies would easily pass into the 
belief in the existence of one i ' ~ 

naturally attribute to spirits tJ 
vengeance or simplest fonn of justice, and t 
which they themselves feel. The Fu^ans appear to be in this 
respectin an intermediate condition, for when the surgeon on board 
the "Beagle" shot some young ducklings as specimens, York 
Minster declared in the most solemn manner, " Oh, Mr. Bynoe, 
" much rain, much snow, blow much;" and this was evidently 
a retributive punishment for wasting human food. So again be 
related how, when his brother killed a " wild man," storms long 
raged, much rain and snow fell. Tet we could never discover 
that the ruegiana believed in what we should call a Grod, or 
practised any religious rites; and Jemmy Button, with justitiable 
pride, stoutly maintained that there was no devil in his land. 
This latter assertion is the more remarkable, aa with savages the 
belief ia bad spirits is far more common than that in good 

The feeling of religious devotion is a highly complex one, 
consisting of love, complete submission to an eialted and 
mysterious superior, a strong sense of dependence," fear, 
reverence, gratitude, hope for the future, and perhaps other 
elements. No being could experience so comples an emotion 

able article on ths 
meats of Raligion.' by 
Pike, ID ' AnthTOpolct^H 
I, 1870, p. Ixiil. 


tj6 The Descent of Man. Taht 1. 

nalil advanced in bis intellectual and moral faculties to at leaat 
a moderately high level. Nevertheless, we see some distant 
approach to this state of mind in the djep love of a di^ for liis 
master, associated with complete submission, some fear, and 
. perhaps other feelings. The behaviour of a d(^ when returning 
to his master after an absence, and, as 1 may add, of a monkey 
to his beloved keeper, is widely different from that towards their 
fellows. In the latter case Uie transports of joy appear to bo 
somewhatless.and the sense of equality is shewn in every action. 
Professor Braubach goes so &r as to maintain that a d(% looks 
on his master aa on a god." 

The same high mental facnltiea which first led man to believe 
in unseen spiritual agencies, then in fetishism, polytheism, and 
ultimately in monotheism, wonld infallibly lead him, as long as 
his reasoning powers remained poorly developed, to various 
strange superstitions and customa Many of these are terrible 
to think of— such as the sacrifice of human beings to a blood- 
loving god; the trial of innocent persons by the ordeal of poison 
or fire; witehoraft, Ac. — yet it is well occasionally to reflect on 
these superstitions, for they shew us what an infinite debt of 
gratitude we owe to the improvement of onr reason, to science, 
and to our accumulated knowledge. As Sir J. Lubbock" has well 
observed, " it is not too much to saj that the horrible dread of 
" unknown evil hangs like a thick cloud over savage life, and 
" embitters every pleasure." These miserable and indirect 
consequences of otu highest faculties may be compared with the 
incidental and occasional mistakes of the instincts of the lower 

" 'Religion, Moral, &c.. der Dar- 

" 'Prehisi 

toric Ttmeis' 3nd edIL 

win'.chen Art>Leh™.' 1869, ^ 53. 

p. 571. Ji 

L this work (p. 571) 

[t is said (I>r. W. Lander Liadsay, 

there will 1 

le found an eieelleat 

'JcBrna! of Mental Sgieac*,' [871, 

BCOOUDt of 1 

the mmy tirao^e iinil 

p. 43), that Baoon long ago, and the 

p« . nnmi, held the same notioiu 



The moral sense — FuBdamental proposition — The qualities of social 
snimals — Origin of socisbilitj' — Strui^le between opposed instiflots — 
Man a sodal animal — The more endnring social instincts conqner otlier 
less persistent instincts — The social virtues alone regarded by savages — 
The self-reEanling virtBcs acquired at a later stage of dcTclopment— 
The importance of the judgment of the members of the same community 
on coniluct — Transmission of moral tendenuies — Snmmary. 

I FULLY subscribe to the judgment oE those writers' who] 
inaJiitiiJii that of all the differencee between maa and the 
lower animals, the moral sense or conscience is by far thej 
Diost important. This sense, as MacMntosh' remarks, " has ai 
" rightful supremacy over every other principle of human 
"action;" it is summed up in that short but imperious word 
mtgJd, so full of high significance. It is the most noble of all 
the attributes of man, leading h'TH without a moment's hesita- 
tion to risk his life for that of a fellow-creature ; or after due 
dehberation, impelled simply by the deep feeling of right or 
duty, to sacrifice it in some great cause. Immanuel Kant 
exclaims, " Duty I Wondrous thought, that workest Eeithei by 
" fond insinuation, flattery, nor by any threat, but merely i^ 
"holding up thy naked law in the soul, and so extorting for 
"thyself always reverence, if not always obedience; before 
" whom all appetites are dumb, however secretly they rebel ; 
" whence thy original ?" ' 

This great question has been discussed by many writers* of 
consummate ability ; and my sole excuse for touching on it, is 
the impossibility of here passing it oyer; and because, as far as I 
know, no one has approached it exclusively from the sida of 
natural history. The investigation possesses, also, some in- 

' See, for instance, on this subject, and Moral Siaenoe,' 1868, p. 543- 

Quatrefaees, • Dnitif de l'Esp*ce 725) of twenty-sii British authors 

Hnmaine," 1881, p. 21, &c who have written on this sabject, 

* ' Dissertation on Ethical Philo- and whose names are fiimiliar to 
•ophy,' 1837, p. 231, &e. every I'eader ; to these, Mr. bain'» 

' 'Metaphysics of Etliica,' trans- own name, and those of Mr. Leekj, 

lated I? J. W. Semple, Edinbarg h, Mr. Shadworth Hodgson, Sir J. 

;83B, p. 138. Lubbock, and others, might 1m 

• Mr. Saia gives a list (' Mental added. 


The Descent of Man. 


t interestj as au attempt to see how far tho stTnly of 
the lower animals throws light on one of the highest psjchical 
Acuities of man. 
I Tho following proposition seems to me in a higli degree 
I probable — namely, that any animal whatever, endowed with 
.' well-marked social instincts," the parental aad filial aifeotiona 
bein^ here included, would inevitably ai;c[uiie a moral sense or 
conscience, as soon as its intellectual powers had become aawell, 
, or nearly as well developed, as in man. Por.jJrsWj, the social 
Instincts lead an animal to take pleasure in tjie society of its 
fellows, to feel a certain amount of sympathy with tJiKm, and to 
perform various services for them. The services may be of a 
definite and evidently instinctive natui'e ; or there may bo only 
a wish and readiness, as with most of the h^her social aDimalS; 
to aid their fellows in certain general ways. But these feelings 
and services ai'e by no means extended to all the individnals of 
the samespecies, only to those of the same association. Secondly, 
as Doon as the mental fecultles had become highly developed, 
im^es of all past actions and motives would be incessantly 
paaaing thiongh the brain of each individual ; and that feeling 
of di'^satisfaction, or even misery, which invariably results, as we 
shall hereafter see, from any unsatisfied instinct, would arise, 
as often as jt was perceived that the enduring and always 
present social mstinct had yielded to some other instinct, at the 
time stronger, but neither enduring in its nature, nor leaving 

" Sir E Brodie, aftei obseriing 

all this, he also remi 

u-ks, '■ii;as is 

that moD IS a socint ouiiDal (' ViJ- 

" my own belief, the 

molia feelings 

chologiwl Enquiries,' 1854, p. 192). 

■' are not innate; hut 

acquired, they 

" are not for that rea 

isoa less natu- 

"not thi> to settle the disputed 

" ral." It is with he 

sitation that I 

" question as to the eiistencc of a 

all from so 

" moral sense r- Similar ideas hnre 

profound a thinker, 

. but it can 

probnblj occurred l« many persons. 

hardly he disputed that the social 

as thev did long ago to Marcos 

feelines are insttnctir< 

9 or mnate in 

Aureliiu. Mr. J, S. MUl speaks, in 

the lo«er animals; ai 

sd why should 
in? Mr. Bdn 

they not be so in m* 

ism,' (1864, pp. 45,46).of thesuoial 

(see, for instance, 'Th. 

■ Emotions and 

feelin]js as a "powerfnl natural 

the Will,' 1865, p. 48 

I) and others 

" sentiment," and as "the nataral 

belisTe that the moiv 

" basis of sentiment for utiiitariaB 

quired by each indi 

lidual during 

" moralitj." Again he sajs, " Ijke 

hislilfetime. On the 

general theory 

" til* other acquired capacities above 

of evolution this Is 

« least ei- 

tremely improbable. 

The ignoring 

"not a part of our nature, is 

of all transmitted m. 

mtal qualities 

"a natural ont-growth fTom it; 

), bo hereafter 

" capable, like them, in a certain 

jndged as a most serio 

■us bkitiish ia 

IJ snwtl degree of apringing up apon- 

the works of Mr. Mil; 


Ohap. it. Moral Sense. 99 

bnhind it a very vhid impression. It is clear that manj in- 
Btinotive desires, SBoh as that of hunger, are in fheir natnre of 
ehort dAiratiou; and after being eatisfled, are not readily or 
j vividly lecallsd. Thirdly, after the power of language had been 
1 acquired, and the wishes of the commanitj could be expressed, 
>|the commoa opinion how each member ought to act for the 
ipublio good, would naturally become in a paramount degree 
the guide to action. But it should be home in mind that how- 
ever great weight we may attribute to public opinion, our regard 
for the approbation and disapprobation of our fellows depends 
on sympathy, which, as wo shall see, forma an essential j)art of 
the social instinot, and is indeed its foundation-stone, Lastly, 
habit in the individual would ultimately play a very important 
part in guiding the conduct of each member ; for the social in- 
stinct, tc^ther with sympathy, is, like any other instinct, greatly 
strengthened by habit, and so consequently would be obedience 
to the wishes and judgment of the community. These seyeral 
subordinate propositions must now be discussed, and some of 
them at considerable length. 

I It may be well first to premise that I do not wish to maintain 
that any strictly social animal, if its intellectual fstcultjes were 
to become as active and as highly developed as in man, would 
acquire eiaetiy the same moral sense as ours. In the same 
manner as various animals have some sense of beauty, thongh 
they admire widely different objects, so they might have a aensr^ 
of right and wrong, thongh led by it to follow widely different 
lines of conduct. If, for instance, to take an extreme ease, men 
were reared under precisely the same conditions as hive-bees, 
there can hardly be a doubt that our unmarried females would, 
like the worker-bees, think it a sacred duty to kill their brothers, 
and mothers would strive to kill their fertile daughters ; and no 
' one would think of interfering." Nevertheless, the heo, or any 

* Mr. H. Sidgv/ick remai'ks, in 
an able discussion on this suljject 

(the 'Academy,' JniiB 15th, 1872, eame illustiHtion, sajs, the prijt- 

(p. 331), " a superior bee, wa may ciples of social duty would be thus 

" ftiel sure, would iitpjre to a milder rfv«rsed ; and by this, 1 preBQme, 

■' solution of the population ques- she nieana that the fulfilment of a 

" tion." Judging, however, from socia,! duty would tend to the injury 

the habits of muny or most savi^es, of individuals ; but she overlooks 

man BoWes th^ problem by female the fact, which ahe would doubtless 

infanticide, po.yandry and promia- admit, that the instiucts of the bee 

cuoDs intetcouTse ; therefore it may have been acqulrcil for the good of 

w«ll be doubted whether it would the community. She goes so far aa 

be by s milder method. Kiss tr, say that If the theory of ethjca 


Tlie Descent of Man. ranr I. 

other social animal, would gain in our siippo 
appears to me, pome feeling of right or wrong, o 
For each indiTidnal would Iiave an inward sense of possessing 
certain stronger or more endnring instincts, and others less 
strong or enduring ; so tliat there would often be ft straggle es to 
which impulse should be followed; and satisfnction, dissatis- 
faction, or evon misery would be felt, as past impressions were 
compared during tlieir incessant passage through the mind. In 
this case an inward raonitor would tell the animal that it would 
have been better to haTe followed the one impulse rather than 
the other. The one course or^ht to have been followed, and the 
other ought not; the one would have been right and the other 
wrong ; but to these terms I shall recur. 

Socmbility. — Animals of many kinds are social ; we find even 
distinct species living together; for example, some American 
monkcTS ; and united flocks of rooks, jackdaws, and starUnge 
Man shews the same feeling in hisstrong love for the dog, which 
the Aog returns with interest. Ererj one must have noticed ho* 
miserable horses, dogs, sheep, &c., are when separated from 
their companions, and what stiijng mutual affection the two 
former kinds, at least, shew oa their reunion. It is curious to 
speculate on the feelings of a dog, who will rest peacefully for 
hours in a room with his master or any of the family, withoiul 
the least notice being taken of him ; but if left for a short time 
by himself, barks or howls dismally. We will confine our 
attention to the higher social animals ; and pass over insect^ 
nithough some of these are social, and aid one another in manj 
important ways. Tlifi. mnst f""''"''" nint'.lfj_L"rTJri] Jn ther^ 
hi gher animals is to w a rn one anoth mLfif dargs r by mp priB dU^ 
th e uni ted sens es of all. Every sportsman knows, as Dr. Jaeger 
remarks,' how difficult it is io approach animals in a herd or 
trooji. Wild horses and cattle do not, 1 believe, make anj 
danger-sign al^: but the attitude of any one of them who first 
discovers an enemy, warns the otherB. Babbits stamp loudly on 
the ground with their "hmS^feet as a signal : sheop and iihamois 
do the same with their forefeet, nttering likewise a nhistlo. 


generally jlcoep!«d, " I cannot but 
" believe that in the hour of their 
" trinmph would be sounded the 
" kuell of the Tittue of roaQkiodl" 
It is to U hoped that the belief in 
i>j pflrnianeDce of virtue on this 

earth is not held by many pet 
on BO Beak a tenure. 

' ' Die Darwin'sche Tbeorie, 

■'Mr. B. Brown in' Pr-c.Zo. 
Soc' IS8S, p. 400, 


of a troop of monkeys acts as tlie sentinel, and titters cneH 
expressive both of danger and of safety.* So cial animftls perform 
m any littlo Bervices for each o ther : horses mbble,- and cowa lick 
each other, on any spot which itoies: monkeys senreh eaoh 
other for eternal parasites ; and Brehm states that after a, troop g> 
of tlie Cfrcopitkneus griseo-viridis has rushed through a thomy 
brake, each monkey stretches itself on a braach, and another 
monkey sitting by, "conscientiously" examines its fur, and 
extracts OTory thorn or burr. 

A nimals also rende r moreJmggrtajrLge^ces to one a noth er : 
thus wolves and some other beas ts of prey hunt in ^ckSj^andg-j 
aid" one another in attacking tJie ir yictims. Pelicans fish ia 
concerts The" Hamadrjas baboons turn over stones to find 
insects, &o. ; and when they come to a large one, as many as can 
stand round, turn it over together and share the booty. SooiaL 
a njmalsmutiiallvfieffiTid each othe r. Ball bisons in N. America^ ®? 
when there is danger, drive the cows and calves into the middle 
of the herd, whilst they defend the outside. I shall also in a 
future chapter give an account of two young wild bulls at 
Chillingtiani attacking an old one in concert, and of two stallions 
together trying to drive away a tidrd stallion from a troop of 
mares. In Abyssinia, Brehm encountered a groat tvoop of 
baboons, who were crossing a valley : some had already ascended 
the opposite monatain, and some were still in the valley : the 
latter were attacked hy the dogs, but the old males immediately 
hurried down from the rocks, and with mouths widely opened, 
roared so fearfully, that the dogs quickly di'cw back. They 
were again encouraged to the attack; but by this time all tha ' 
baTxwns had reasccoded the heights, excepting a young one, 
about six months old, who, loudly calling for aid, climbed on a 
block of rock, and wb'I surrounded. Now one of the largest ( 
males, a true hero, came down again from the mountain, slowly 
wont to the young one, coaxed him, and triumphantly led him ' 
away — the dogs being too much astonished to roako an attack, ; 
I cannot resist giving another scene wtiioh was witnessed by this 
same naturalist ; an eagle seized a young Cercopithecus, which, 
by clinging to a branch, was not at once carried off; it cried / 
.oudlyfor assistance, upon which the other members of the troop, 
with much uproar, rushed to the rescue, surrounded the eagle, 

! Brehm, ' Thierlebon,' B. i. 1864, the evi lenoo of Altarei, whose ob- 
s, 52, 79. For the case of tho Een-tuiusBrehm thinksouiii' triui- 
monkeya estrauting ihnrpit iVnm wm-thu Pivr iht, 4—^,, ^f iha nl.» 
each other, see s. 54. 
to the HamftdTyas tnrnini 
stones, lie fact ia gii-en (s. 


The Descent of Man. 

ftiid pulled out BO many featliers, that he no longer thought 
of his prey, but only how to escape. This eagle, as Brehm 
ro marka. aasTirc dl 

is certain that associated animals have a fcelii^ of love for 
eMh other, which is not felt by non-social adult animals. How 
far in most cases they actually BympatMse in the paine and 
pleasures of others, is more doubtful, especially ■with respect to 
pleasures. Mr. Buxton, however, who had excellent means of 
observation," states that hismacana, ■which hved free in Norfolk, 
took " an extravagant interest " in a pair witli a nest ; and when- 
ever the female left it, she was surronnded by a troop " scream- 
" ing horrible acclamations in her honour." It is often difficult 
to Judge whether animals have any feeling for the sufferings of 
others (tf their kind. "Who can say what cows feel, when they 
surround and stare intently on a dying or dead companion; ap- 
parently, however, hb Hoazeau remarks, they feel no pity. Ttwtt 
animals sometimes are far from feeling any sympathy is too 
certain ; for they will espel a wounded animal from the herd, or 
gore or worry it to death. This is almost the blackest fact in 
natural history, unless, indeed, the explanation which has been 
suggested is true, that their instinct or reason leads them to 
expel an injured companion, lest beasts of prey, including man, 
should be tempted to follow the troop. In this case their con* 
duct is not much worse than that of the Forth American Indiana, 
who leave their feeble comrades to perish on the plains; or the 
Fijians, who, when their parents get old, or fall ill, bury thom 

Many animals, however, certainly sympathise with each other's 
distress or danger. This is the case even with birds. Capt 
Stansburj ^ found on a salt lake in Utah an old and completely 
blind pelican, which was very fat, and must have been well fed 
for a long time by his companions. Mr. Blyth, as he informs 

and Mag. of Hat. Hist.,' 

'• Mr. Belt gives the case of 

gua, which was heard soreamliig; fc 
nearv wo ^ul^ in j^^ orest. aD 

1- Kov 

= by i( 

The i 

r J. Lubbock, ' Prehistori.J 
imes," 2nd edit. p. 44S. 
" Ab quotei by Mr. L. H. Morgan 

niained fee* to fioe ; a: 

uralist in Nicsragua,' 1874, 


Chap. IV. Moral Seme. IC3 

me, saw Indian crows feeding two or three of their companions 
which were blind ; and I hare huard of an analogous case with 

3 cock. We may, if we choose, call these actions 
instinctiTS ; but such casee are much too rare for the develop- 
ment of anj special instinct." I have myself seen a dog, who 
never passed a cat who lay sick in a basket, and was a great 
friend of his, withont giving her a few licks with his tongue, the 
surest sign of kind feeling in a dog. 

"^ It must be called sympathy that leads a courageous dog to 
fly at any ond who strikes his master, a-s ho certainly will. I 
saw a person pretending to beat a lady, who had a very timid 
little d(^ on her lap, and the trial had never been made before , 
thelittle creature instantly jumped away, but after the pretended 
beating was over, it was really pathetic to see how perseveringly 
he tried to lick bis mistress's face, and comfort her. Brehm " 
states that when a baboon in confinement was pursued to be 
punished, the others tried to protect him. It must have been 
sympathy in the cases above given which led the baboons and 
Cercopiiheci to defend their young comrades from the dogs and 
the eagle. I will give only one other instance of sympathetic 
and heroic conduct, in the case of a Uttle American monkey. 
Several years ago a keeper at the Zoolc^cal Gardens shewed me 
some deep and scarcely healed wounds on the nape of his own neck, 
inflicted on him, whilst kneeling on the floor, by a fierce baboon. 
The little American, monkey, who was a warm friend of this 
keeper, lived in the same large compartment, and was dreadfully 
afraid of the great baboon. Neverthelefs, as soon as he saw his 
friend In peril, he nished to the rescno, and by screams and bites 
so distracted the baboon that the man was able to escape, after, 
as the surgeon thought, running great risk of his life. 

Besides love and sympathy, animals eihibit other qualities 
connected with the social instincts, which in us would be called 
moral ; and I agree with Agassiz ^' that di^s possess somethii^ 
very like a conscience. 

Dogs possess some power of selt-command, and this does not 
appear to be wholly the result of fear. As Branbach" remarks, 
they will refrain from stealing food in the absence of their 
master. They have long been accepted as the very type oi 
fidelity and obedience. But the elephant is likewise very faith- 
ful to his driver or keeper, and probably considers him as the 

" aid tg a Enffever spring! from sj 
**pathy proper:" 'M«ital andMc 
Science,- !86S, p, 245. 


[04 Tlie Descent of Man. Pabt \. 

leader of the herd. Dr. Hooker informs me that an elephant, 
which he waa riding in India, became so deeply Iw^ged that he 
remained stuck fast until the nest day, when he ■was estricated 
hy men with ropes. Under Bach cLrcTimstances elcphanta will 
seize with their trunks any object, dead or alive, to place nnder 
their knees, to prevent their sinking deeper in the mnd ; and the 
driver was dreadfully afraid lest the animal should have seized 
Dr. Hooker and crushed bi ni to death. But the driver himself, 
as Dr. Hooker was assured, ran no risk. This forbearance under 
an emergency so dreadful for a heavy animal, is a. wonderful 
proof of noble fidelity.^* 

All nnimiils livinp; in ah ody.ffhic h def end themse lves or attack 
, th eir enemies in c onc ert, must indeed be in some degree Sithful 
' to o ne another; and those that" follow aTeaileFmuST^ En some 
lie^Se^obedieiit. When the baboons in Abyssinia" plunder n 
garden, they silentl j follow their leader ; and if an imprudent 
yoiu^ animal maJces a noise, he receives a slap from the others 
to teach him silence and obedience. Mr. Galton, who lias had 
eicelleat opportunitieB for observing the half-wild cattle in S. 
Africa, says,^ that they cannot endure even a momentary separa- 
tion from the herd. Th ^ are eEsentially slavish, and accept th» 
com mon determination, "seekJ cg-no^'EeHCTlor tKaiTfFbe led bj 
any one oic who ha s enough "sel l-reliajcfe. to .accept the position. 
The men who"l)feak in theseanimals for haicess, watch assidu- 
ously for those who, by grazing apart, shew a self-reliant dis- 
position, and these they train as fore-oxen. Mr, Galton adds 
that such am"mals are rare and valuable ; and if many were bom 
they would soon be eliminated, as lions ajre always on the look- 
out for the individuals which wander fcom the herd. 

■With respect to the impulse which leads certain annuals to 
associate together, and to aid one another in many ways, we 
may infer Uiat in most eases they are impelled by the same 
sense of satiEfacfion or pleasure which they experience in per- 
forming other instinctive actions; or by the same sense of 
dissatiBfaetion as when other instinctive actions are checked. 
We see this in innumerable instances, and it is illustrated in a 
striking manner by the acquired instincts of our domesticated 
animals; thus a young shepherd-d(^ del^hts in driving and 
running round a flock of sheep, but not in worrying them ; a. 
young fos-hoimd delights in hunting a fox, whilst some other 
kinds of dogs, as I have witnessed, utterly disregard foxes. What 


' Himalayan " Ste I, 

' Thlcrloben,' B, i 

linper ■ 


tMAV. IV. 

jilorat Heuse. 


B strong feolbg of inwsijrd satisfaction must impel a bird, so full 
of activity, to brood day after day oTer her eggs. Migratorji 
birds aro quite miserablo if stopped from migrating; perlia]!s 
they caijoy starting on their long flight ; but it is hard to believe 
that the poor pinioned goose, described by Audubon, wliich 
started on foot at the proper time for its journey of probably 
more than a thousand miies, could have felt any joy in doing so. 
Some instincts are determined solely by painful feelings, as fc; 
fear, wtiich leads to self-preservation, and is in some cases directed 
towards special enemies. No one, I presume, can analyse the 
sensations of pleasure or pain. In many instances, however, ii 
is probable that instincts are persistently followed from the 
mere force of inheritanoo, without the stimulus of either 
pleasure or pain, A young pointer, when it first scents game, 
apparently cannot help pointing. A squirrel in a cage who pats 
tie nuts which it cannot eat, as if lo bury them in the ground, 
can hardly he thought to act thus, either from pleasure or pain. 
Hence the common assumption that men must be impelled to 
every action by esijeriencing some pleasure or pain may bo erro- 
neooa. Although a habit may be blindly and implicitly 
followed, independently of any pleasure or pain felt at the 
moment, yet if it be forcibly and abruptly checked, a vague 
Bense of dissatisfaction is generally experienced. ' 

It lias often been assumed that animals were in the first place 
rendered social, and that they feel as a consequence uneomfort^ 
able when separated from each other, and comfortable whilst 
together ; but it is a more probable view that these sensations 
were first developed, in order til"*'- ♦■''"'^ jVniTT"'! " which would 
profit by living in society, should be induced to live togethe r, 
in the saiiie manner as the sense of-firraglTana^fLe pleasure o\ 
eating were, no doubt, first acquired in order to induce animals 
to eat. The feeling of pleasure from society is probably an/ 
extension of the parental or filial affections, since the social! 
instinct seems to be developed by the young remaining for a I 
long time with their parents; an d this extension may be at tri- \ 
buted in uart to hab it, but chie^y to nat ural s election. . With_J 

tho inStviduals w M^_t ook the greatest p leasur e in society ^ 
woni dnicst oscaua^various dangers^ whil at. t.lnwi t,b»t i-areil 
least for their comrades, and lived ^o lit aiy, would p erish in 
graitetlfiuioberB,. With respect to tho origin of the paSStal 
nnd filial affections, which apparently lie at the base of the 
social instincts, we know not tho steps by which they liave 
bsea gained; but we may infer that it has been to a large 
axifiiit through natural selection. So it Las almost certainly 


'Hie Descent of Man. 

been with the unusual and opposite feeling of hatred between 
the ceareBt relations, as with tho workor-bees which kill their 
brothcr-droceSj and with the queen-bees which kill their 
daughter-queens ; the desire to destroy their nearest relations 
having be^ in this case of serrice to the community. Parental 
affection, or some feeling which replaces it, has been developed 
in certain aninjals extremely low in the scale, for example, in 
star-fishea and spiders. It is also occasionally present in a few 
members alone in a whole group of animals, as in the genus 
Porficula, or earwigs. 

The all-important emotion of sympathy is distinct from that 
of love. A mother may passionately lovo her sleeping and 
passive infant, but she can hardly at such times be said to feel 
sympathy for it. The love of a maa for his dog is distinct from 
sympathy, and so is that of a dog for Ms master. Adam Smith 
formerly argued, as has Mr. Bain recently, that the basis of 
sympathy lies in our strong retentiveness of former states of 
pain or pleasure. Hence, " the sight of another person en- 
" during hunger, cold, fatigue, revives in us some recollection of 
" these states, which are painful even in idea." We aie thus 
impelled to relieve the sufferings of another, in order that our 
own painful feelings may be at tho same time relieved. In like 
manner we are led to participate in the pleasures of others.^' 
But I cannot see how this view explains the fact that sympathy 
is excited, in an immeasurably stronger degree, by a beloved, 
than by an indifCerent person. The mere sight of suffering, 
independently of love, wotUd snfflce to call up in ns vivid 
iwollections and associations. The explanation may lie in the 
fact that, with all animals, sympathy is directed solely towards 
the members of the same community, and therefore towards 
known, and more or lees beloved members, but not to all tho 
individuals of the same species. This feet is not more sur- 
prising than Oiat the fears of many animals should be directed 
against special enemies. Species which axe not social, such as 
lions and tigers, no doubt feel sympathy for the suffering of 
their own young, but not for that of any other animal. With 

*' See the first and striking " or others in his stead, maj make 

chapter ia Ad^^m Smith's 'Theoiy ^' up, by sympathy and good ofHces 

of Moral Sentiments.' Also Mr. " returned, for all the sacrifioe." 

Bain's 'Mental and Moral Science,' But If, aa appears to be the ease, 

1868, p. 2*4, and 375-282. Mr. sympathy is sti-ictly an instinirt, 

Itain states, tliat "sympathy is, in- its eieroise woald give dire.^t plea- 

' jire<tlif, a source of pleasure lo sure, in the same manner as th« 


CBiip. IV. Moral Sense. 107 

mankuid, eelfisliceas, esperienco, and imitation, probably odd, 
as Mr. Bain has ehewn, to the power of Eympathy ; for we aro o-oy 
l ed by the hope of receiTiag p[ood in retiirn to perform act8_ ^ Wecl 
oT" sympathotic kindness to others; and sympathr^is muc h 
t rtjeaKthcncd by habi t. In however complex a raanner thi3~N 
feetii^ may have originated, as it is one of high importance to [ 
all those animals which aid and defend one another, it will havo I 
been increased through natnial seiection ; for those commu- i 
nities, which included tJie greatest nnnjber of the most sympa- ( 
thetic members, woidd flourish best, and roar the greatest \ 
niinibor of offspring. ■-— ) 

It is, however, impossible to decide in many cases whether 
certain social instincts have beun acquired through natural 
selection, or are the indirect result of other instincts and 
foculties, such as sympathy, loflson, experience, and a tendency 
to imitation ; or again, whetlier they are simply the result of 
long .continued habit. So remarkable an instinct as the placing 
sentinels to warn the community of danger, can hardly have 
been the indirect result of any of these feculties ; it must, tbore- 
fore, have been directly acquired. On the other hand, the habit 
followed by tho males of some soc jal HiiimplK of ilpfpni^ in} ; \h n 
community, and of attacking their RnfimipH nr t>ipir pri-y in 
concert, may~perhap8 have originated from mu tual sympa thy ; 
but eou rafie, anci m rnost cases strength, miist have b een 
previously acquired, probably through naturaFselec tion. 

Of the various instincts and habits, some are much stronger 
than others ; that is, some either give more pleasure in their 
performance, and more distress in their prevention, than others ; 
or, which is probably quite as important, they are, through 
inheritance,, more persistently foEowed, without exciting any 
special feeling of pleasure or pain. We are ourselves conscious 
that some habits are much more difGcnlt to cure or change than 
otliers. Hence a struggle may often be observed in animals 
between different instincts, or between an instinct and some 
habitual disposition ; as when a dog rushes after a hare, is 
rebuked, pauses, hesilatfes, pursues again, or returns ashamed lo 
his master ; or as between the love of a. female dog for her young 
puppies and for her master,— for she may be seen to slink away 
to them, as if half ashamed of not accompanying her master. 
But the most curious instance known to me of one instinct 
getting the better of another, is the migratory instinct conquer- 
ing the maternal instinct. The former is wonderfully strong; a 
confined bird will at the proper season beat her breast against 
the wires of her cage, until it is bare and bloody. It causes 
young salmon to leap oot of the fi;esh water, in which they could 


loS ■ The Descent of Man. Past I. 

coatiaue to exist, and tiius imintentionally to commit suioido. 
Every one knows how strong the maternal instinct is, leading 
even timid birds to fiice great danger, though with hesitation, 
and in opposition to theinstinct of self-preseryation. Neverthe- 
less, the migratory instinct is so powerful, that Me in the autumn 
Bwfjlows, hoosa-martins, and swifts frequently desert their 
tender young, leaying them to perish miserably in their nests." 

We can perceive that an instinctive impulse, if it be in any 
way more beneficial to a species than some other or opposed 
instinct, would be rendered the more potent of tlio two through 
natural selection ; for the individuals which hod it most strongly 
developed would survive in larger numbers. Whether this is the 
case with the migratory in comparison with the maternal instinct, 
mf.y be doubted. The great persistence, or steady action of the 
former at certain seasons of tiio year during the whole day, laK^ 
gi e t ibr a time paramount force. 

M n a somal anima!. — Every one will admit that man is a 
so al being. We see this in his dislike of solitude, and in his 
wish fo society beyond that of his own family. Solitary con- 
fin m nt is one of the severest punishments which can be 
TTifli ted Some authors suppose that man primevally lived in 
singl families ; but at the present day, though singlo families, 
o only two or three together, roam the soHtudes of some savage 
land they always, as far as I can discover, hold friendly 
relat ns with other families inhabiting the same district. Such 
famih occasionally meet in council, and unite for their common 
d f noe. It is no argument against savage man being a social 
anunal that the tribes inhabiting adjacent districts are almost 
al ay at war with each other ; for tiis social instincts nevei 

t nd t all the individuals of the same species. Judging from 
tl e a lalogy of the majority of the Quadrumana, it is probable 
th t t! early ape-like progenitors of man were likewise social ; 
but this s not of much importance for us. Although man, as 

" This lact, the Eev. I. Jenyns hatched. USaay birds, not jet old 

•taU9 (see his edition of '^ White's enongli for s prolonged flight, ai-e 

Kat. Hist, of Selborne,' 1853, p. likewise deseitod and left behind. 

204) «as first recorded by the illu:^ See Blackwall, 'Researches in 2oo- 

Irions Jenner, in 'PhiL Transact.' logy,' 1834, pp. 108, 118. For same 

182*, and has since been confirmed additional evidence, although this 

by several observers, especially by is not wanted, aea Leroy, 'Letti'eii 

Mr. Ulackwalt. This latter carefnl Phil.' 1802, p. 217, For Swifts, 

Dbserver ei.imined, late in the Gould's 'Introduction U> the Bii'^ia 

auCuion, duiing two yeai^s, thirty- of Great Britain,' 1823, p. 5. Simi- 

til nests { he found that twelve lar cases have been observed in 

coBtaioed young dead birds, live Canada by Mr. Adams; 'Pup. 

contained eggs on the point of being Science Rpview,' July 187J, [l 

hEtcheU, and three, eggs not nearly 'JS'i. 


Chap, IV, Moral Sense. icxj 

hs now exists, has few special instincts, taving lo3t aiij whicli 
his early progenitors may have possessed, this is no reason why 
he should not have retained from an extremely remote period 
Bome degree of instinctiTe love and sympathy for his fellows. 
We are indeed all conscious that we do possess such sympathetic 
foelingB ; '^ but our consciousness does not tell ns whether they 
are instinctive, having originated long ago in the same manner 
as with the lower animals, or whether they have been acquired 
IS during our early years. A s man is a social anim al , 

for these qualities are common to most social animals, lie woitld 
consequently possess some capacity for self-command. He 
would from an i nherited tendency be willing to defend, in 
conc ert with oth eig^his fellow-mcn "I~an d woiii3~tie reai^to'aid .}^ 
them m any way, which did n ot too greatly inteijere with^ his 
own weliare or his own stro n g 'desi res. 

The social animals which stand afthe bottom of the scale are I 
guided almost esclusively, and those which eland higher in the ! 
scale are largely guided, by special instincts in the aid which 
they give to the members of the same community; but they are | 
likewise in part impelled by mutual love and sympathy, assisted • 
apparently by some amount of reason. Although man, as just 
remarked, has no special instincts to tell him how to aid his 
fellow-men, he still has the impulse, and with his improved 
intellectual faculties would naturally be much guided in this 
respect by reason and experience. Instinctive sympathy would 
also cause him to value highly the approbation of his fellows ; 
for, as Mr. Bain has clearly shewn,^ the love of praise and 
the strong feeling of glory, and the still stroller horror of scorn 
and infamy, " are due to the workings of sympathy." Conse- 
quently man would be influenced in the highest degree by the 
wishes, approbation, and blame of his fellow-men, as eipre^ed 
by their gestures and language. Thus the social instincte, 
which most have been acquired by man in a very rude state, 
and probably even by his early ape-like progenitors, still give 
the impulse to some of his best actions ; but his actions are in a 
higher degree determined by the expressed wishes and Judgment 

» Hnme rem^tke ('Ad Enquiry 


Concerning the Pnnciplss of Morals,' 

edit, of XTSl, p. 132), "ThBre seems 

"the latt. 

'■ s necessity fcr lonfessing that the 

"choly d: 

- different to us, but that the view 


no The Descent of Man. i'abt I. 

or his fellow-men, and nnfortTinatelj Tory oftea by his own strong 
Belfish. desires. But as love, sympathy and self-command become 
Btrengthened by habit, and as the power of reasoning becomes 
dearer, so that man can value Justly tLe judgments of his 
foUows, he will feel himself impelled, apart from any transitory 
pleasure or pain, to certain lines of conduct. He might then 
deulare— not that any barbarian or uncultdvaled man could 
thus think — I am the supreme judge of my own conduct, and ii; 
tho words of Kant, I will not in my own person Tiolat« the 
dignity of humanity. 

Tiie more endariay Social Instincts conqver the lts» persisleiil 
lastincti.—'We have not, however, as yet conadered the main 
point, on which, from our present point of view, the whole 
question of tho moral sense turns. Why should a man feci that 
be ought to obey one instinctive desire rather than another ? 
Why is he bitterly regretful, if ho has yielded to a strong sense 
of self-preseryation, and has not risked his life to save that of a 
fellow-creature ? or why does he r^ret having stolen food from 

I t. is evident in t haJraJplitce. that wiih.mankindjhe instinc- 
ti ve impu l ses hav e different d^rees of strength ; a savage will 
ris k his own life to save thatataJSsnber of the same community, 
but will be w holly indifferent about a stranger : a young and 
timid mother urged by the maternal instinct will, without a 
moment's hesitation, run the greatest danger for her own infant, 
but not for a mere fellow-creature. Nevertheless many a 
civilized man, or even boy, who never before risked his life for 
another, but full of courage and sympathy, has diregarded 
the instinct of self-preservation, and plunged at once into a 
torrent to save a drowning man, though a stranger. In this case 
man is impelled by the same instinctive niotise, which made the 
heroic little American monkey, formerly described, save his 
keeper, by attacking the great and dreaded baboon. Such 
actions as the above appear to be the simple result of the greater 
strength of the social or maternal instincts than that of any 
other instinct or motive; for they are performed too instan- 
taneously for reflection, or for pleasure or pain to bo felt at the 
time ; though, if prevented by any cause, distress or even misery 
might be felt. In a timid man, on tlie other hand, the instinct 
of self-preservation might be so strong, that he would be unable 
to force himself to run any such risk, perhaps not even for his 
own child. 

I am aware that some persons maintain that actions perfoimcd 
impnlaively, as in the above cases, do not come under the 
dominion of the moral sense, and cannot bt called moral. Thcj 


Chap. TV. Moral Sense. 1 1 1 

confine this tenn to actions done de!iberaie!y, after a victorj 
OTer opposing desires, or when prompted by some eialted 
motive. But it appears seoreoly possible to draw any clear line 
of distinction of tius kind.^^ Afi fai as exalted motives are 
concerned, many instances have been recorded of savages, 
destitute of any feeling of general benevolence towards mankind 
and not guided by any religious motive, who have dehberateij 
sacrific ed their lives aa prison ers," rather than IJetraj their 
c omrades .; and snrely their conduct ought 16 he 'COnBidefed a«" 
moral As far as deliberation, and the victory over opposing 
motives are concerned, animals may be seen 3oiil)fmg~'Bglween 

from d anger; yet their actions, though done for the goocT of 
others, are not called moral. Moreover, anything performed 
very often by us, will at last be done without deliberation or 
hesitation, and can then hardly be distinguished from an 
instinct ; yet snrely no one will pretend that such an action 
ceases to be moral. On the contrary, we all feel that an act 
cannot be considered as perfect, or as performed in the most 
noble manner, unless it be done impulsively, witliout deliberation 
or offortj in the same manner as by a man in whom the 
qualities are innate. He who is forced to overcome his fear 
want of sympathy before be acts, deserves, however, in one way 
higher credit than the man whose innate disposition leads hjiri 
to a good act without effort. As we cannot distinguish between 
motives, we rank all actions of a certain class as moral, if 
performed by a moral being. A mora] being is one who is 
capable of comparing his past and future actions or motives, and 
of approving or disapproving of them. We have no reason to 
suppose that any of the lower animals have this capacity ; 
therefore, when a Newfoundland dog drags a child out of the I 
water, or a monkey faces danger to rescue its comrade, or takes 1 
chaise of an orphan monkey, we do not call its conduct moral. \ 
But in the case of man, who alone can with certainty be ranked 
as a moral being, actions of a certain class are called moral, 
whether performed deliberately, after a struggle with opposing 

■« I refer here to the distinction " material nnd formal moralil.y !a 

tericd and forvml morality. I am " tinctions." 

giadtofindthatFr(.f.Hiiilej('Cri- "" 1 hsve eiien one encli caw, 

tiqnes and Addresses,' 1873, p. 287) namely of three Patagonian Indians 

lakes th« same view on this snbject who preferred being shot, one aft«r 

HB I do. Mr. Leslie SWphen re- the other, to betraying the |)lnn« of 

marto (' Essays on Freethinking and their companions in war (' Jonn-iJ 

Plain Speaking:, 1873, p. 83), " the of Researdes,' IS+S. p. lOS). 
*- metaphysical distinction tetweeu 


1 1 2 The Descent of Man. Past L 

motives, or impulsively through insticot, or &om the offecta of 
slowly-gainod liabit, 

Bnt to retura io our more immediate subject. Although 
Bome instincts are more powerful than others, aud thus lead to cor- 
responding aotioBS, yet it is -untenable, that in man the social 
instincts (including the iove of praise and fear of blame) possess 
greater strength, or have, through long habit, 8cc[uired greater 
strength than the instincts of self-preservation, hunger, Inat, 
Tengeance, &c Why then does man regret, even though trjing 
to banish such regret, that he has followed the one natural 
impulse rather than the other ; and why does he further feel 
that he ought to regret his conduct'/ Man in this respect differs 
profoundly from the lower animals, Nevertheless we can, I think, 
eee with some degree of clearness the reason of this difference. 

Man, from the actii-ity of his mental faculties, cannot avoid 
reflection : past impressions and images are incessantly and 
clearly passing through his mind. Now with those animals 
which live permanently in a body, the social instincts are ever 
present and persistent. Such animals are always ready to uttei 
the danger-signal, to defend the community, and to give aid to 
their fellows in accordance with their habits; they feel at all 
times, without the stimulus of any special passion or desire, 
some degree of love and sympathy for them ; they are unhappy 
if long separated from them, and always happy to be again in 
their company. So it is with ourselves. Even when we are 
quite alone, how often do we think with pleiBure or pain of 
what others think of us, — of their imagined approbation oi 
disapprobation ; and this all follows from sympathy, a funda- 
mental element of the social instincts. A man who possessed 
no trace of such instincts would be an unnatural monster. On 
the other hand, the desire to satisfy hunger, or any passion such 
as vengeance, is in its nature temporajy, and con for a time be 
fully satisfied. Nor is it easy, perhaps hardly possible, to call 
up with complete vividness the feeUng, for instance, of hunger ; 
nor indeed, as has often been remarked, of any suffering. The 
instinct of self-preservation is not felt except in the presence of 
danger; and many a coward has thoi^ht himself brave until ho 
has met his enemy (hoe to face. The wish for another man's 
property is perhaps as persistent a desire as any that can lie 
named; hut even in this case the satisfaction of actual pou- 
session is generally a weaker feeling than the desire ; many a 
thief, if not a habitual ona, after success has wondered why ho 
stole some article.*' 

" Enmity or hatred seems also perhaps inars to than na^ othr-i 
to be 3 highly persistent fteiing, that can V« Mmed. &vv is At- 


CHip. IV. Moral Sense. 1 1 3 

A mail cannot prevent past impressions often repassing thtoae'i 
his inind ; he will tJius be driven to make a comparison between 
the impresBiona of past liimger, vengeance satisfied, or danger 
shnnned at other men's cost, with the almost eTor-present 
instinct of Bjmpathy, and with his earlj knowledge of what 
others condder as praiseworthy or blameable. This knowledge 
cannot be banished from his miud, and from instmctivesjmpathy 
is esteemed of great moment He will then feel as if he had 
been baulked in following a present instinct or habit, and this 
■with all animals causes dissatisfaction, or even misery. 

The aboTe case of the swallow affords an illnstiatjon, though 
of a reversed natnre, of a temporary thoagh for the time strongly 
peisistcnt instinct conquering another instinct, which is usually 
dominant over all others. At the proper season these birds 
seem all day Jong to be impressed with the desire to migrate ; 
their habits change ; they become restless, are noisy, and con- 
gregate in flocks. Whilst the mother-bird is feediag, or brooding 
over her nestlings, the maternal instinct is probably stronger 
than the migratory ; but the instinct which is the more jwrsis- 
tent gains the victory, and at last, at a moment when her young 
ones are not in sight, she takes flight and deserts them. When 
arrived at the end of her long journey, and the migratorj 
instinct has ceased to act, what an i^ony of remorse the bird 
would feel, if, from being endowed with great mental activity, 
she could not prevent tlie image constantly passing through her 
mind, of her young ones perishing in the bleak north from cold 
and hunger. 

fined as hatred of another for some had done him an injuij and had 
eioallenca or inecMBs; and Bacon become hia enemy. Nor is it pri>- 
ioiiEtB (Essay ix.),"OF all other Ubie that the primitive conscience 
'* affectiotis envy is the most im- nould reproach a man for injuring 
" pcrtone and ccntinnaL" Dogs are his enemy ; rather It would re- 
very apt to hate both strange men pioaeh him, if he bad not revenged 
ud dugs, especially if they himself. To do good in return fur 

h m family, tribe, or clan; 

of molality to which it ma 

r be 

e« g would thus seem to be 

doubted whether the aoeial inat 

d 3 certainly a most per- 

would, by themselves, have eve 


n It seems to be the 

ua. It ia necessary that the& 

9 in- 

p m and converse of the 

Btincts, togetJier with ayicipj 


SOL instinct. From *h;it 

should ha™ been highly cullii 


h avagee, it wenld appear 

and eitended by the aid of re 


t something of the same. kind 

instruction, and the love or fe 

la good with them. If this be 

God, before any such golden 


it would be a small step in 

would ever bo thought of 


one to transfer sncli feelings to 


member of the same iribe if he 


1 14 Tfie Descent of Man. Pab~ 

At tho moment of action, man will no donbt be apt to foUoM 
the stronger impuiae; aad though this may occasionallj 
prompt him to tte noblest deeds, it will more commonly leail 
him to gratify his own desires at the expense of other men. 
But after their gratification, whea past and weaker impresBiona 
are judged by tho ever-enduring sociol instinct, and by his deep 
regard for the good opinion of his fellows, retribution will surely 
come. He will then feel remorse, repentance, regret, or shame 
this latter feeling, however, relates almost eiclusively to the 
judgment of others. He will consequently resolve more or less 
firmly to act difforeutly for fho future ; and this is conscience ; for 
conscience looks backwards, and serves as a guide for the future. 

The nature and strength of the feelings which we call regret, 
shame, repentance or remorse, depend apparently not only on 
' the strength of the violated instinct, but partly on the strength 
of the temptation, and often still more on tho judgment of 
■ onr fellows. How far each man values the appreciation of 
others, depends on the strength of his innate or acquired 
feeling of sympathy; and ou his own capacity for reasoning out 
the remote consequences of his acts. Another element is most 
important, although not necessary, the reverence or fear of the 
Gods, or Spirits believed in hy each man : and this applies 
especially in cases of remorse. Several critica have objected 
that though some slight regret or repentance may be eiplaine"! 
by the view advocated in this chapter it is impossible thus to 
account for the soul-shaking feeling of remorse But I can see 
little force in this objeetion. My cntios d) not define what 
they mean by remorse, and I can find no definition implying 
more than an overwhelraing sense of reppntanco Eemor^e 
seems to bear the same relation to repentance, as rage does to 
anger, or agony to pain. It is far from strange that an instinct 
t«i strong and so generally admired, as maternal lore, should, if 
disobeyed, lead to the deepest misery, as soon as the impression 
of the past cause of disobedience is weakened. Even when an 
action is opposed to no special instinct, merely to know that our 
friends and equals despise ns for it is enough to cause great 
misery. Who con doubt that the refusal to fight a duel through 
fear has caused many men an agony of shame ? Many a Hindoo, 
(t ia said, has been stirred to the bottom of his soul by having 
Irfirtaken of unclean food. Here is another case of what must, 1 
think, be called remorse. Dr. Landor acted as a magistrate in 
West Australia, and relates,^ that a native on his fitrra, after 
losing one of his wives from disease, came and said that " he was 

" 'Insanity in r.cliitioo to Law;' Ontaiio, United States, 1S7'., p. ii. 


CoAP. IV. JIforai Sense. 1 1 5 

" going to a distant tribe to spear a woman, to satiafy liie Beam 
" of duty to his Mrife. I told him that if he did bo, I would 
" send him to prison for life. He remained about the farm for 
" Bom3 montlis, hut got exceedingly thin, and complained that 
" he could not rest or eat, that his wife's spirit was haunting 
" him, becauBe he had not taken a lifo for hers. I was in- 
" exorable, and assured him that nothing should save him if ha 
■' did." NevertJielees the man disappeared for more than a year, 
and then returned in high condition ; and his other wife told 
Dr. Landor that her husband had taken the life of a woman 
belonging to a distant tribe; but it was impossible to obtain 
legal evidence of the act. The breach of a rule held sacred by 
tlie tribe, will thus, as it seems, give rise to the deepest focliugs, 
—and this qxdte apart from tbe social instincts, excepting in so 
far as the rule is grounded on the judgment of the community. 
How so many strange superstitions have arisen throughout the 
world wo know not ; nor can we tell how some real and great 
crimes, such as incest, havo come to be held in an. abhorrence 
(which is not however quit* universal) by the lowest savages. It . .. 
is even doubtful whether in some tribes incest would be looked on 
with greater horror, than would the marriage of a man with a 
woman bearing the same name, though not a relation. '■ To 
violate this law is a crime which the Australians hold in the 
greatest abhorroneo, in this agreeing esaetly with certain 
tribes of North America. When the question is put in either 
district, is it worse to kill a girl of a foreign tribe, or to marry 
a girl of one's own, an answer just opposite to ours would be 
given without hesitation."™ We may, therefore, reject the 
belief, lately insisted on by some writers, that the abhorrence of 
incest is due to our possessing a special God-implanted con- 
science. On the whole it is intelligible, that a man utged by 
BO powerful a sentiment as remorse, though arising ss above 
explained, should bo led to aet in a minner, nhich ho has been 
taught to believe serves as an expiation, such as delivering 
himself up to justice. 

Man prompted by his conscience, wdl tlirou^h long habit \ 
acquire such perfect self-command, that his de^rcs and passions I 
will at last yield instantly and without a struggle to his social i 
sympathies and instinctSjincluding his feeling for the judgment of \ 
hisl'ellows. The stillhungry,orthe still revet^fiil man willnot '| 
think of stealing food, or of wreaking his vengeance. It is possible i 
or ns we shall hereafter see, even probable, that the habit of self- j 
command may, like other habits, be inherited. Thus at last man i 

» E. a Tylor in ' Contemporarjr R*vi«w,' April, 1?73, p, 707. 


! i6 Tlw Descent of Man. Pabt I 

comes to fel, through acquired and perhaps inhorited habit, that 
it is best for him to obey his moro peisistent impulses. The 
imperious word o'tnU seems merely to imply tho consciousness of 
the existence of a rule of conduct, however it may have 
originated. Formerly it must haTe been often vehemently 
urged that an insulted geutleman ought to flght a duel. "We 
oveu Bay that a pointer ougM to point, and a retriever t« 
retrieve game. If they fail to do so, they fail in their duty 
and act wrongly. 

If any desire or instinct leading to an action opposed to the 
good of others still appears, when recalled to mind, aa strong 
as, or stronger than, the social instinct, a man will feel no keen 
regret at having followed it ; but he will be conscious that if his 
conduct were known to his fellows, it would meet with their 
disapprobation ; and few are so destitute of symjBthy as not to 
feel discomfort when this is realised, If he has no such 
sympathy, and if his desires leading to bad actions are at tho 
time strong, and when recalled are not over-mastered by the 
persistent social instincts, and the judgment of others, then ha 
is essentially a bad man;* and the sole restraining motive loft 
is the fear of punishment, and the conviction that in the long 
run it would be best for his own selfish interests to regard the 
good of others rather than his own. 

It ia obvious that every one may with an easy coasoienee 
gratify his own desires, if they do not interfere with his social 
instincts, that is with the good of others ; but in order to be quite 
free from self-reproach, or at least of ansiety, it is almost neces- 
eary for him to avoid the disapprobation, whether reasonable or 
not, of his fellow-men. Nor must he break through the fixed 
habits of his life, especially if these are supported by reason ; 
for if he does, he will assuredly feel dissatisfaction. He must 
likewise avoid the reprobation of the one God or gods in whom, 
according to his knowledge or superstition, he may believe ; but 
in this case the additional fear of divine punishment ofton 

The slrkSy Social Virlrtes at first akme regarded. — The above 
view of the origin and nature of the moral sense, which tella na 
what we ought to do, and of the conscience which reproves us if 
we disobey it, accords well with what we see of the early and 
nndeveJoped condition of this faculty in mankind. The virtues 
which must be practised, at least gencraOy, by rude men, so 

•■ Dr. Prosper DesfiDc, Id his msay curiona esses of the wurst 
' Vsjchologie Katurelle,' 1863 (torn. ciiminBls, who app»renlly havfl been 


Ucir. 1 '.'. Moral Sense. 117 

that they may associ.vte in a body, are those which are etiU 
recognised as the most important. But_th£y_axajauetiBed 
almost e sclpsiTely ia relation to the men of the same tribe; and 
their oppoai tes are not regarded as crimes in relation to the men 
of othe r tribes. No tribe could hold t<^ether if murder, 
robbery, treachery, &a., were common ; consequently snch 
erimcB within the limits of the Barae tribe "are branded with 
" evei'lasting inE^my;"" but excite no such sentiment beyond 
these limits. A North-American Indian is well pleased with 
himself, and is honoured by others, when he scalps a man of 
another tribe ; and a Byak cuts off the head of an unoffending 
person, and dries it as a trophy. The murder of infants hss 
preyailed on the lai^est scale throughout the world,** and has 
met with no reproach ; but infanticide, especially of females, has 
been thought to be good for the tribe, or at least not injurious. 
Suicide during former times was not generally considered as a 
crime,^ but rather, from the courage displayed, as an honourable 
act ; and it is still practised by some semi-ciTilised and savage 
nations without reproach, for it does not obyiously concern 
others of the tribe. It has been recorded that an Indian Thug 
conscientiously regretted that he had not robbed and strangled 
as many trav^ors as did his father before him. In a rude state 
of civilisation the robbery of strangers is, indeed, generally 
considered as honourable. 

Slavery, although in some ways beneficial during ancient 
times," ia a great crime ; yet it was not so regarded until quite 
recently, even by the most civihzed nations. And this was 
esxMcially the case, because the slaves belonged in general to a 
race different from that of their masters. As barbarians do not 
regard the opinion of their women, wives are commonly treated 
like slaves. Most savages are utterly indifferent to the sufferings 
even delight in witnessing them. It is well 

'1 See m tble article in the 

of Eoropean Morals,' vol. i. 1869, 

North British RetIb™,' 1807, p. 

p. 223. With respect to Bav^es, 

895. Se« also Mr. W. Bagehof> 

Mr. Winnood Reade inforuis me 

arUclM on the Importance of Obe- 

that the negroes of West Africa 

dieace and Coherence to Primitire 

often commit suicide. It is well 

Man, in the 'Fortnightly Beview,' 

13fi7, p. 539, and 1S68, p. 457, Ac 

the miserable aborlglnea of South 

" The fullest noconnt which I 

America, after the Spanish conquest. 

tsve met with is hy Dr. Gerltud, in 

For New Zealand, se* the voyage oi 

his 'Oeber dan AusBterben der 

the "Novara," and for the Aleutian 

Nalorviilker,* 1868; but 1 shall 

Islands, Muller, aa quoted by Hon- 

have to recur tfl the enfaiect of 

zean, • Les Faeult^s Mentales," &c. 

infanticide In H futnre chapter. 

torn. ii. p. 136. 

» See Mr. Bagehot, <Plv«os and 

tioc C3 Splclde in Leetj'e ' HUtory 

Folitua,' 1872, p. 73. 


t\8 The Descent ?J Man. w 

knowii tJmt the women and children of the North-Aj 
Indians aided in torturing their enomios. Some saTages take a 
horrid pleasure in cruelty to ani-tiais,^ and humanity Is an 
unknown virtue. NeyettheleBa, besides the family atfections, 
kindness is common, especially during sickness, between the 
members of the same tribe, and is sometimes extended beyond 
these limits. Mungo Park's touching account of the kindness of 
the negro women of the inferior to him is well known. Many 
instances could be giyon of the noble fidelity of sawiges towards 
each other, but not to strangers ; common experience justifies 
the masim of the Spaniard, "Never, never trust an Indian." 
There cannot be fidelity without truth ; and this fundamental 
Tiituo is not rare between the members of the same tribe ; thiiB 
Mimgo Park heard the n^ro women teaching their young 
children to love the truth. This, again, is one of the Turtuea 
which bocomeB so deeply rooted in the mind, that it is sometimea 
practised by sav^ea, even at a high cost, towards strangers ; but 
to lie to your enemy has rarely been thought a sin, as the history 
of modem diplomacy too plainly shews. As soon as a tribe has 
a recognised leader, disobedience becomes a crime, and even 
abject submission is looked at as a sacred virtue. 

As durii^ rude times no man can bo useful or faithful to liia 
tribe without courage, this quaUty has universally been plaeed 
in the highest rank; and although in civilised countries a 
good yet timid man may be far more useful to the community 
than a brave one, we cannot help instinctively honouring the 
latter above a coward, however benevolent. Prudence, on tha 
other hand, which does not oonoem the welfare of others, though 
a very useful virtue, has never been highly esteemed. As nc 
man can practise the virtues necessary for the welfare of his 
tribe without self-sacriflce, self-command, and the power ot 
endurance, these qualities have been at all times highly and 
most justly valued. The American savage voluntarily submits 
to the most horrid tortures without a groan, to prove and 
strengthen his fortitude and courage; and we cannot help 
admiring him, or even an Indian I^kir, who, from a foolish 
religious motive, swings suspended by a hook buried in his 

The other so called self-regarding virtues, which do not 
obviously, though they may really, affect the welfare of the tribe, 
have never been esteemed by savages, though now highly 
appreciated by civilised nations. The greatest intemperanoa 

" See, for instance, Mr. Hamilton's account of tlie KalfirB, ' Anthro- 
pnlcgiial Review,' J 870, p. it. 


CHiP. IT, Moral Seme. 1 19 

is no reproach, ■with ssTages. Utter licentiousness, and un- 
natural crimes, preTail t« an astounding estent.^ As soon, 
howeTer, as itiarriago, whether polygamous, or rooaogamous, 
becomes common, jealousy will lead to the inculcation of female 
virtue; and this, being honoured, will tend to spread to the 
unmarried females. How slowly it spreads to the male sex, 
we see at the present daj Chastity eminently requires self- 
command; therefore it has been honoured from a very early 
period in the moral history of oiTiljsed man. As a consequence 
of this, the senseless practice of cehbocy has been ranlied from a 
remote period as a virtue." The hatred of indecency, which 
appears to us so natural as to bo thought innate, and which is 
BO valuable an aid to chastity, is a modem virtue, appertaining 
eiclusiYely, as Sir G. Staunton remarks," to civilised life. This 
is shown by the ancient religious rites of various nations, by the 
drawings on the walls of Pompeii, and by the practices of many 

We have now see n t hat a otioi^ are regard^ by s avages , and 
we re probably so re garded 67 piiraevaT man, as good^r^^d, 
sol ely as t hey obviously affect the weliSeof thalMEie, — not that 
of_^e_specieSj_not that of an individual memberof the. trilM!, 
This conclusion agrees well with the belief that the so^alled 
mora! sense is aboriginally derived from the social instincts, for 
both relate at first exclusively to the community. The chief 
causes of the low morality of savages, as judged by our 
standard, are, firstly, the confinement of sympathy to the same 
tribe. Secondly, powers of reasoning insufficient to recognise 
the bearing of many virtues, especially of the self-regarding 
virtues, on the general welfare of the tiibe. Savages, for 
instance, fail to trace the multiplied evils consequent on a 
want of temperance, chastity, &c. And, thirdly, weak power 
of self-command; for this power has not been strengthened 
through long-continned, perhaps inherited, habit, instruction and 

I have entered into the above details on the immorality of 
savages," because some authors have recently taken a high view 
of their moral nature, or have attributed most of their crimes to 
mistaken benevolence.*' These authors appear to rest their 

" Mr. M'Lennaa lias given " ' Embassy to Chioa,' vol. iL d. 

('Primitive Marriage,' 1865, p. 343. 

176) » good coliectioii of f.cta OB " See on this Bnbjeet copious 

this bsarl, evideneo in Cliap. vii. of Sir J. Lnb- 

« Letliy, 'Histoiy of European bock, 'Origiu of Civilisalioa.' 1870. 

McisIe,' tol. 1. 18G9, p. 109. " For instance I.«oty, 'Hiat 
Eoropean M(ral», vul. u \^. 131. 


Tke Descent of Man 

cfinelusion on savages possessing those Tirtues which aro sor- 
viceable, or even necessary, for the existence of the family and of 
the tribe,— qualities which they undonhtedly do possess, and often 
in a high di^gree. 

CottdvAing Ifemar^s. — It was assumed formerly by philosophers 
of the derivatiye" school of morals that the foundation of morality 
lay in a form of Selfishness; but more recently the "Greatest 
"happiness prineipla " has been brought prominently forward. 
It is, however, more correct to speak of the latter principle as 
the standard, and not as the motive of conduct Nevertheless, all 
the authors whose works I have consulted, with a few escep- 
liuns,*' write as if there must be a distinct motive for every 
action, and that this must be associated with some pleasure or 
displeasure. But man seems often to act impulsively, that is 
from instinct or long habit, without any consciousness of pleasure, 
in the same manner as does probably a bee or ant, when it 
blindly follows its instincts. Under ciremnstacces of extreme 
peril, as durii^ a lire, when a man endeavours to save a fellow- 
creatara without a moment's hesitation, he can hardly feel 
pleasure ; and still less has he time to reflect on the dissatisfaction 
which he might subsequently esperience if he did not make the 
attempt. Should he afterwards reflect over his own conduct, he 
would feel that there lies within him an impulsive power widely 
different from a search after pleasure or happiness; and this 
seems to be the deeply planted social instinct. 

In the case of the lower animals it seems much more appro- 
priate to speak of their social instincts, as having been developed 

" This term is nsed in sn able 

" nesa eitra-regarding impntse, di- 

nrtide in the ' Westminater Review," 

" reeled tavatis something that is 

Oct. 1869, p. 493. For the " Greatest 

" not pleasure; that in many cases 

" bsppiness prineiple," see J. S. Mill, 

" with the self-reEaidiug that the 

" Will rewgoises ('Syslem of 

" tivo do not easily coexist in the 

Lngio,' vol. iL, p. 422) in the clearest 

manner, that actiona may bo per- 

A dim feeling that onr impolses do 

formed thmugh habit withont the 

not by any maaos always arise from 

Sidgwick alBO. in his Essay on 

pleasuve, lias, 1 cannot bnt think, 

Vleasnre and Desire ('The Con- 

biipn one chief cause of the accept- 

ance of the intulUve theory of 

C71), remarks: "To snm np, in 

morality, nod of the rejection of the 

■' coattaTMitlon of the doctrine that 

utilitarian or " Greatest happiDcss " 

theory. With respect to the latter 

" nlisaya directed towards the pro- 

theory, the standard and the motive 

of conduct have no doubt often beet 

" unrselves, 1 woqW mdintain that 

confused, bnt fhev are really i: 

" KB fiuJ eiferywhire ic consoions. 

u>me degree blended. 


OKiP. IT Moral Sense. 121 

for tho general good ratter than for the general happiiiees of ths 
Bgecies. The term, general good, may be defined as the rearing 
of tho greatest number of individnals in tall Tigonr and healtli, 
with all their faeuitics perfect, under the conditions to which 
they are subjected. As the Rocial instineta both of man and tho 
lower aiijrmils have no doubt been developed by nearly tho wuno 
steps, H; would be advisable, if found practicable, to use the 
enine definition in both cases, and to take as tho standard of 
morality, the general good or welfare of the community, rather 
than the general hax>piness ; but this definition would perhaps 
require some limitation on account of political ethics. 

When a man risks his life to save that of a fellow-croatTire, it 
seems also more correct to say that he acts for the general good, 
rather than for the general happiness of mahkind. No douht 
the welfare and tho happiness of the individual uanaJly coincide ; 
and a contented, happy tribe will flourish better than one that 
is discontented and unhappy. We have seen that even at an 
early period in the history of man, the expressed wishes of tho 
community will have naturally influenced to a largo extent the 
conduct of each member ; and as all wish for happiness, the 
" greatest happiness principle " will have become a most im- 
portant secondary guide and object ; the social instinct, however, 
together vfith sympathy (which leads to omr regarding the 
approbation and disapprobation of others), having served as tho 
primary impulse and guide. Thus the reproach is removed of 
laying the foundation of the noblest part of our nature in the 
base principle of selfishness; unless, indeed, the satisfaction 
which every animal feels, when it follows its proper instincts, 
and the disaatisfaction felt when prevented, be called selfish. 

The wishes and opinions of the members of the same community, 
expressed at first orally, but later by writing also, either form 
the sole guides of our conduct, or greatly reinforce tho social 
instincts ; such opinions, however, have sometimes a tendency 
directly opposed to these instincts. This latter fact is well 
exemplified by the Lav) of Honour, that is, the law of the opinion 
of our equals, and not of all our countrymen. The breach of 
this law, even when the breach is known to be strictly accordant 
with true morality, has caused many a man more agony than a 
real crime. We rect^uise tlie same influence in the burning 
Bense of shame which most of us have feit, even after the interval 
of years, when calling to mind some accidental breach of a 
trifling, though fixed, rule of etiquette. The judgment of the 
comrnumty will generally be guided by some mde experience of 
what is best in the long run for all the members ; but this ju(^- 
moot will not rarely eir fwm ignorance and we;ik powers of 


122 Ths Descent of Man. Part 1, 

reasoning. Hence the strangest customs and superstitions, in 
complete opposition to the true welfare and happiness of man- 
kind, have become all-powerful througliout tho world. We see 
this in the horror felt by a Hindoo who breaks his caal«, and 
in many other such cases. It would be difBcult to distinguish 
between the remorse felt by a Hindoo who has jieldud to the 
temptation of eating unclean food, from that felt after committing 
a thoft ; but tho former would probably be the more severe. 

How so many absurd rules of conduct, as well as so majo- 
absurd reUgious beliefs, have originated, we do not know ; noi 
how it is that they have become, in all quarters of the world, so 
deeply impressed on the mind of men; but it is worthy of 
remarkthat a belief constantly inculcated during the early years 
of life, whilst the brain is impressible, appears to acquire almost 
the nature of an mstinct ; and the very essence of an instinct is 
that it is followed independently of reason. Neither can we say 
why certain admirable Tirtues, such as the love of truth, are 
much more highly appreciated by some savage tribes than by 
others;" nor.again, why similar differences prevail even amongst 
highly civilised nations. Knowing how firmly fixed many 
strange customs and superstitions have become, we need fee) no 
surprise that the self-regarding virtues, supported as they are by 
reason, should now appear to us so natural as to be thought 
innate, although they were not valued by man in his earlj- 

Notwithstanding many sources of doubt, man can generally 
and readily distinguish between the higher and lower moral 
rules. The higher are founded on the social instincts, and relate 
to the welfare of others. They are supported by the approbation 
of our fellow-men and by reason. The lower rules, tliough some 
of them when implying self-sacrifice hardly deserve to be called 
lower, relate chiefly to self, and arise from public opinion, ma^ 
tnred by experience and cultivation; for they are not practised 
by rude tribes. 

As maa advances in civilisation, and small tribes are united 
.nto larger communities, the simplest reason would toll each 
individual that he ought to extend his social instincts and 
sympathies 'x> all the members of tho same nation, though 
jiersonally unknown to him. This point bemg onoo reached, 
there IE only an artificial barrier to prevent his sympathies 
eitendmg to the men of all nations and races. If. indeed, such 
men are separated from him by great differences in appearance 

" GiBd instances ar* civen by in his 'ContribuHoDS to th" Th«,i-. 
S«^ t5, If 


Ceap. IV, Moral Sense. 123 

or habits, es[wrienpe unfortunately siiewB us how long it is, 
before wo look at them as our fellow-creaturea. Sympathy 
beyond the confines of m^, that isj humanity to the lower 
animals seenis to be one of the latest moral acquisitions. It is 
apparetUy unfelt by savages, except towards their pets. How 
little the old Eomans knew of it is shown by their abhorrent 
gljtdiator/ai eshihitions. The very idea of hnmanity, as lar as I 
could observe, was new to most of the Gnuchos of the Pampas. 
This virtue, one of the noblest with which man is endowed, 
seems to arise incidentally from our sympathies becoming more 
tender and more widely diffused, until they are extended to all 
sentient beings. As soon as this virtue is hononted and practised 
by some few men, it spreads through instruction and example 
to the jouog, and eventually becomes incorporated in public 

The highest possible st^e in moral culture is when we re- 
c(^nise that we ought to control our thoughts, and " not even in 
" inmost thought to think again the sins that made the past so 
" pleasant to us."" 'Whatever makes any bad action familiar to 
tite mind, renders its performance by so much the easier. As 
Marcus Aurelius long ago said, "Such as aro thy habitual 
" thoughts, such also will be the character of thy mind ; for the 
" soul is dyed by the thoi^hts."" 

Our great philosopher, Herbert Spencer, has recently explained 
his views on the moral sense. He says,*' " I believe that the 
" experiences of utility organised and consolidated through all 
" past generations of the human race, have been prodncing 
" corresponding modifications, which, by continued transmission 
" and accumulation, have become in us certain faculties of 
" moral intuition — certain emotions respondii^ t« right and 
" wrong conduct, which have no apparent basis in the individual 
" experiences of ntihty." There is not the least inherent 
improbability, as it seems to me, in virtuous tendencies being 
more or leas strongly inherited ; for, not to mention the various 
dispositions and habits transmitted by many of our domestic 
animals to their oftspring, I have heard of authentic cases in 
which a desire to steal and a tendency to lie appeared to run 
in families of the upper ranks; and as stoahi^ is a rare crime i « 
the wealthy classes, we can hardly account by accidental coinci- 
dence for the tendency occurring in two or three membeis of 

" Teanrsnn, ' Idylls of the King," Aurolins waa born A.D. 121. 

n, 244. " Lettiii to Mr. Mill in Itain'i 

« 'The Thoughts of the Emperor 'Mental and Moral Science," 1B68 

H Aorelius Anioninna,' Eng. trans- p. 723. 
Ut.,SDdeilit.,lS60,p.lt3. M&rcu 


124 The Descent of Man. Fast L 

the same family. If bad tendencies are transmitted, it is pro- 
bable that good ones are litewise transmitted- That tlie statn 
of the body by affecting tie braiii, has great iniiiienco on tlia 
moral tendencies is known to most of those who havo suffertd 
from chronic derangements of the digestion or liver. The Eamo 
fact is liliewiae Bhewn by tho " perversiori or destruction of the 
" moral sense being often one of the earliest symptoms of mental 
"derangement^"' and insanity is notoriously oft«n inherited. 
Escept through the principle of the transmission of moral ten- 
dencies, we cannot understand the differences believod to exist in 
this respect between the various races of mankind. 

ETen the partial transmission of virtuous tendencies would 
be an immense assistance to the primary impulse derived directly 
and indirectly from the social instincts. Admitting for a momen*^ 
that viriiuous tendencies are inherited, it appears probable, at 
least in such cases as chastity, temperance, humanity to animals. 
&e,, that they become first impressed on the mental organization 
through habit, instruction and example, oontinned during 
Feveral generations in the same family, and in a quit* subordinate 
degree, or not at all, by the indiyiduals possessing such vii-tues 
ha ving R i"'™°^'''i best in the struggle for IHe. K^ chief sourc* 
of doubt with respect to any Buch inJieritance, is that senselasB 
customs, superstitions, and tastes, sach as the honor of a Hindoo 
for unclean food, ought on the same principle to be transmitted. 
I have not met with any evidence in support of tJie transmission 
of superstitious customs or senseless habits, although in itself it 
is perhaps not less probable than that animals should acquii« 
inherited tastes for certain kinds of food or fear of certain foes. 

Finally the social instincts, which no doubt were acquired by 
man as by the lower animals for the good of the community, 
will from the first have given to him some wish to aid his 
fellows, some feeling of sympathy, and have compelled him to 
regard their approbation and disapprobation. Such impulses 
will have served him at a Tery early period as a rude rule of 
right and wtorg. But as man gradnally advanced in intellectual 
power, and was enabled to trace the more remote consequences 
of his actions; as he acquired sufficient knowledge to reject 
baneful customs aad snperstitions ; as he regarded more and 
more, not only the welfare, hut the happiness of his fellow-men ; 
as from habit, following on beneficial experience, instruction 
and example, his sympathies became more fender and widely 
diffcsed, extending to men of all racts, to the imbecile, maimed, 

" Jfandslty, 'Body .ind Mind,' IflTO, p. 00. 


Cflip. IV. Summary. 1 2 5 

will other lossless membeis of society, and finally to the lowei 
animals, — ao would the standard of itis morality rise higher and 
higher. And it is andmittei?' by moralists of the derivative 
sehool and by some iutnitionists, that the standard of morality 
has risen since an early jwriod in the history of man.*" 

As a struggle may sometiiaes be seen goicg on between the 
various instincts of the lower animals, it is not surprising that 
there should be a struggle in man between Hb social instincts, 
with their derived virtues, and his lower, though momentarily 
stronger impulses or desires. This, ss Mr. Gallon" has remarked, 
is all the less surprisiiig, as man has omei^d from a state of 
barbarism within a comparatively recent period. After havirg 
yielded to some temptation we feel a sense of dissatisfaction, 
shame, repentance, or remorse, anali^ous to the feeUngs caused 
by other powerful instincts or desires, when left unsatisfied or 
baulked. We compare the weakened impression of a past 
temptation with the ever prosent social instincts, or witi habits, 
gained in early youth and strengthened during our whole lives, 
until they have become almost as strong as instincts. If with 
the temptation still before las we do not yield, it is because 
either the social instinct or some cusiom is at the moinonl 
predominant, or because we have learnt that it will appear to us 
hereafter the stronger, when compared with the weakened im- 
pression of the temptation , and we reahse that its violation would 
cause us suffering. Looking to future generations, there is no 
cause to fear that the social instinctB will grow weaker, and we 
may expect that Tirtuous habits will grow stronger, becoming 
perhaps fixed by inheritance. In this cose the struggle betweuii 
. our h^her and lower impiUses will bo leas severe, and virtue 
will be triumphant, 

no doubt tliat 
1 and that of 

the highest animal is immense. An anthropomorphous ape, if 
he could take a dispassionate view of his own cstee, would admit 
that though he could form an artfnl plan to plunder a garden — 
though he could use stones for fighting or for breatiug open 
nuls, yet that the thought of fashioning a stone into a tool was 

*" A writer ia the ' North British 
Review" (Julj 1869. p. 531), well 

meat, eipre^ses himself Ettouglv id 
favour of this coDcluiion. Ur. 
Lecky ('Hist, of Morals,' voi. i. p. 

" See his cemarkable work i 


1 26 T//e Descent of Man. I'Aft'P 

IwViia iiojond his scope. Still less, a£ lie would admit, could he 
loUow out a ti-ain of metaphysical i-easouing, or solve a mathe- 
matical problem, or reflect on G(S, or admlve a grand aaturul 
scene. Some apes, however, would prohahly declare that they 
could and did admire the beauty of the coloured skin and fur of 
their partners in marriage. 'Jhey would admit, that though they 
coald make other apes understand by cries some of their per- 
ceptions and simpler wants, the notion of expressing definite ideas 
hy definite sounds had never ci'ossed their minds. They might 
insist that they were ready to aid their follow-apes of the same 
troop in many ways, to risk their lives for them, and to take 
charge of their orphans; but they would be forced to acknoiv- 
iedge that disinterested love for all living creatures, the mosl 
noMe attribute of man, was qidte beyond their comprehenKion. 

Kovertheless the difference in mind between man and the 
higher animals, great as it is, certainly is one of degree and not 
of kind. We have seen that the senses and intuitions, the 
various emotions and feculties, such as love, memory, attention, 
curiosity, imitation, reason, &c., of which man boasts, may bo 
found in en incipient, or even sometimes in a well-developed 
condition, in the lower animals. They are also capable of some 
inherited improvement, as we see in tie domestic dog compared 
with the wolf or jackal If it could be proved that certain high 
mental powers, such as the formation of genera! concepts, self-oon- 
Bciousness, &c., were absolutely pecuhar to man, which seems 
rtxtremely doubtful, it is not improbable that these qualities are 
merely the incidental results of other highly-advanced intel- 
lectual faculties; and these again mainly the result of the 
contiuned use of a perfect language. At what age does the 
new-born infant possess the power of abstraction, or become 
self-conscious, and reflect on its own existence? We cannot 
answer ; nor can we answer in regard to the ascending organic 
scale. The hali-art, half-instinct of language still btars Uio 
stamp of its gradual evolution. The ennobling belief in God is 
not universal with man; and the belief in spiritual agencies 
naturally follows from other mental powers. The moral sense 
perhaps aflbrds the best andhighest distinction between man and 
the lower animals ; but I need say nothing on this head, as I 
have so lately endeavoured to shew that the social instincts, — 
the prime principle of mac's moral constitution "'—with the aid 
of active intellectual powers and the effects of habit, naturally lead 
to the golden rule, " Ae ye would that men should do to you, do 
je to them likewise ;" and this lies at the foundation of morality 

•■ ' Tho ThoDght! of Marcos Anreliiis,' &i!., p, 13S, 



tellectual Faculties. 12? 

In the next cliapter I shaii nrafce some few romarkB on Iha 
probable steps and means by whicb tbe several mental and moral 
(koulties of man have been gradually evolTed. That saoh evolu- 
tion is at least possible, ought not to be denied, for we daily sea 
these faicnltiea developing in every infant ; and wo may trace a 
perfectr gra-latjon from the mind of an ntter idiot, lower tliRn 
that of an animal low in the scale, to the mind of a Newton. 

Oh the Development or tee luTBLLKornAi. and Mon.ii, 


AdvaBoement of the intellectDal powew ttirough natnral (election — 
Importance of imitation— Social and moral [acHl-ics— Thfir develoii- 
luent within the limits of the same trihe— Natuta. selectiou lu ati^<:tiug 
civilued nations—Evidence that civilised natioua were once baibarond. 

Tub subjects to be discussed in this chapter are of the highest 
interest, but are treated by me in an imperfect and fragmentary 
manner. Mr. Wallace, in an admirable paper before referred to,' 
ai^ues that man, after he had partially acquired those intel- 
toctual and moral faculties which distinguish him from the 
lower animals, would have been but little liable to bodily 
modifications through natural selection or any other means. 
Porman is enabled through his mental faculties "to keep with 
" on unchanged body in harmony with the changing nniverse," 
He has great power of adapting his habits to new conditions of 
hfe. He invents weapons, tools, and various stratagems to 
procure food and to defend himself. When he migrates into a 
colder climate he uses clothes, builds sheds, and maics fires; 
and by the aid of fira cooks food otherwise indigestibla Ho 
aids his fellow-men in many ways, and anticipates future events. 
Even at a remote period he praetised some division of labour. 

The lower animals, on the other hand, must have their bodily 
stractura modified in order to survive under greatly changed 
conditions. They must be rendered stronger, or acquire more 
effective teeth or claws, for defence against new enemies; oi' 
they must be reduced in size, so as to escape detection and 
danger. When they migrate into a colder climate, they must 
become clothed with thicker fur, or have their constitutioua 
altered. If they fail to be thus modified, they will coase to 

..^ ' 'Aalhropoljgical Revic^i',' Kay 1864, p. cirtii. 


123 Tke Descent of Man ■ , PamJ, 

The ease, howe^isr, is widt^j different, as fe Wallace lias 
with justice insistod, in relation to the intollectual and moral 
fftomlties of man. These faculties are variable; and we have 
every reason to believe that the variations tend to be inherited. 
Therefore, if they were formerly of high importance to primeval 
nwn and to his ape-like progenitors, they would have befin 
perfected or advanced thiongh natnrij, selection. Of the liigh 
importance of the intellectnol faculties there can bo no doubt, 
for man mainly owes to them his predominant position in the 
world. We can Bee, that in the rudest state of society, the 
individuals who were the most sagacious, who invented and used 
the weapons or traps, and who were best able to defend 
themselves, woold rear the greatest noinber of offspring. The 
tribes, which included the largest number of men thus endowed, 
WQuld increase in number and supplant other tribes. Numbers 
depend primarily on the means of subsistence, and this depends 
partly on the physical nature of the country, but in a much higher 
degree OE the arts which are there practised. As a tribe increases 
and is victorious, it is often still further increased by the ab- 
sorption of other tribes.' The stature and strength of the meti 
of a tribe are likewise of some importance for its success, and 
these depend in part on the nature and amount of the food which 
can be obtained. In Eimipe the men of the Bronze period were 
supplanted by a race more powei-ful, and, judging &om their 
eword-handlcH, with larger hands ;' but; their saccess was pro- 
bably still more due to their superiority in the arts. 

All that we know about saTsges, or may infer from their 
traditions and from, old monuments, the history of which is quite 
forgotten by the present inhabitants, shew tliat from the remotest 
times successful tribes have supplanted other tribes. Eelics ot 
eitiuct or forgotten tribes have been discovered throughout the 
civihsed regions of the earth, on the wild plains of America, and 
on the isolated islands in the Pacific Ocean. At the present day 
civilised nations are everywhere supplanting barbarous nations, 
excepting where the climate opposes a deadly barrier; and thoy 
succeed mainly, though not exclusively, through their arts, which 
are the prod nets of the intellect. It is, therefore, highly probable 
that with mankind the intellectual faculties have been mainly 
and gradually perfected through natural selection ; and this con- 
clusion is sufficient for our purposa Undoubtedly it would 
be interesting to trace the development of each separate faculty 

" After a tim* the mcmbera or 1861, p, 131), that thej are the o 

tribes wnic j ara absorbeJ intg desceiidaafs of the same anoeston 
mother tribe assume, as Sir Henry » Moilot, 'So.'. Vaad. So. Kat 

Miiine remarks (' Ancient Law,' 1860, p. 291. 


Cuip. T. Moral Faculties. 129 

from the state in which, it exists ia tie lower animals to that in 
which it exists in man; but neither my abilitj uor knowledge 
pormits the attempt. 

It deserves notice that, as soon as the progenitors of man 
became social (and this prohahlj occnircd at a very early period), 
the principle of imitation, and reason, and experience would 
have increased, and much modifled the intellectuaj powers in a, 
way, of which we see only traces in the lower animals. Apes ate 
much given to imitation, as are tbe lowest savages; and the 
sitople fact previously referred to, that after a time no animal 
can be caught in the same place by the same sort of trap, shews 
that animals leam by experience, and imitate the caution of 
others. Now, if some one man in a tribe, more sagacious than 
the others, invented a new snare or weapon, or other means of 
attack or defence, the plameet self-interest, without the assistance 
of much reasoning power, would prompt the other members to 
imitate him; and all would thus profit. The habitual practice 
of each new art must likewise in some slight degree strengthen 
the intellect. If the new invention were an important one, the 
tribe would increase in number, spread, and supplant other 
tribes. In a tribe thus rendered more numerous there would 
always be a rather greater chance of the birth of other superior 
and inventive members. If such men left children to inherit 
theit mental superiority, the chance of the birth of stiil mote 
ingenious members ■jrould be somewhat better, and in a very 
small tribe decidedly better. Even if they left no children, the 
tribe would stiU include their blood- relations; and it has been 
ascertained by agriculturists* that by preserving and breeding 
from the family of an animal, wtiioh when slaughtered was 
found to be valuable, the desired character has been obtained^ 

Timiiag now to the social and moral faculties. In order that 
primeval men, or the ape-hke pregcnitors of man, should become 
social, they must have acquired the same instinctive feelings, 
which impel other animals to live in a body ; and they no doubt 
exhibited the same general disposition. They would have felt 
uneasy when separated from their comrades, for whom they 
would have felt some degree of love ; they would have warned 
each other of danger, and have given mutual aid in attack or 
defence. All this implies some degree of sympathy, fidelity, and 
courage. Such soci^ qualities, the paramount importanco of 
which to the lower animals is disputed by no one, were no doubt 

* 1 have glvea instances in my ' V.iriilion of Aninala unjcr DoiiimH™- 


130 The Descent of Man. Pabt I. 

acquired by tho progenitors of man in a similar monner, namely, 
through natural selection, aided by inherited habit. When two 
tribes of primeTal man, liring in the same country, caroe into 
competition, if (other circumstances being equal) the one tribe 
included a great number of courageous, symjmthetic and faithful 
members, who were always ready to warn each other of danger, 
to aid and defend each other, this tribe would succeed better and 
conijuer the other. Let it be home in mind how all-important 
in the nerer-ceasing wars of savages, fidelity and courage must 
be. The advantage which disciplined soldiers haYe over undis- 
ciplined hordes follows chiefly from the confidence which each 
man feels in his comrades. Obedience, as Mr. Bagehot has well 
shown,' is of the highest value, for any form of government is 
better than none. Selflsh and contentious people will not cohere, 
and without coherence nothing can be effected. A tribe rich in 
the above qualities would spread and be victorious over other 
tribes : hut in the course of time it would, ju^ing from all past 
history, be in its turn overcomo by some other tribe still more 
highly endowed. Thus the social and moral qualitiea would 
tend slowly to advance and he diffused throughout the world. 

But it may he asked, how within the limits of the same tribe 
did a large number of members first become endowed with those 
social aad moral qualities, and how was the standard of ex- 
cellence raised ? It ia extremely doubtful whether the offepring 
of the more sympathetic and benevolent parents, or of those 
who were tho most faithful to their comrades, would be reared 
in greater numbers than the children of selflsh and treacherous 
parents beloi^ing to the same tribe. He who was ready to 
BBcrifiee his life, as many a savage has been, rather than betray 
hia comrades, woidd often leave no offepring to inherit his noble 
nature. The bravest men, who were always willing to come to 
the front in war, and who freely risked their lives for others, 
would on an average perish in larger numbers than other men. 
Therefore it hardly seems probable, that the number of men 
gifted with such virtues, or that the standard of their excellence, 
could be increased through natural selectioa, that is, by the 
survival of the fittest ; for we are not here speaking of one tribe 
being victorious over another. 

Although the circumstances, leading to an increase in the 
number of thoee thus endowed within the same tribe, are too 
complex to bo clearly followed out, we can trace some of tho 
probable steps. In the first place, as the reasoning powers and 

• Sm a remarkable aeries of arti- April 1,18^; Jnly 1, 1889, siWK 
dn on ' Phjsiis and Politics' ia the separs'tcly jmhliehed. 


Ohaf. V. Moral Facttlties. I JI 

foresight of the membera became improved, eadi man would 
Boon learn that if he aided his fellow-men, he would commonly 
receive aid in return. From this low motdve he might acquiro 
the haWt of aiding his fellows ; and the hahit of performing 
benevolent actions certainly strengthens the feeling of sympathy 
which gives the first impulse to benevolent actions. Habits, 
moreover, followed during many generations probably tend to 
be inherited. 

But another and muct more powerful Btimnlns to the de- 
velopment of the social virtnes, is afforded by the praise and the 
blame of our feliow-men. To the instinct of sympathy, as we have 
already sees, it is primarily doe, that we habitually bestow both 
praise and blame on others, whilst we love the former and dread 
the latter when applied to ourselves ; and this instinct no doubt 
was originally acquired, lite alltbeother social instincts, through 
natural selection. At how early a period the prf^nitors of man 
in the course of tlieir development, became capable of feelingand 
being imjwlled by, the praise or blame of their fellow-creatnres, 
we cannot of course say. But it appears that even dogs appre- 
ciate encouragement, praise, and blame. The rudest savages 
feel the sentiment of glory, as they clearly show by preserving 
the trophiesof their prowess, by their habit of excessive boasting, 
and even by the ertreme care which they take of their per- 
sonal appearance aud decorations ; for tmiess they regarded the 
opinion of their comrades, snob habits would be senseless. 

They certainly feel shame at the breach of some of their leaser 
rules, and apparently remorse, as shewn by the case of the 
Australian who grew thin and could not rest from having 
delayed to murder some other woman, so as to propitiate bis dead 
wife's spirit. Though I liave not met with any other recorded 
case, it is scarcely credible that a savage, who will sacrifice his 
life rather than betray his tribe, or one who will deliver himself 
up as a prisoner rather than break his parole," would not 
feel remorse in his inmost soul, if he had failed in a duty, 
which he held sacred. 

We may therefore conclude that primeval man, at a very 
remote period, was influenced by the praise and blame of bis 
fellows. It is obvious, that the members of the same tribe would 
approve of conduct which appeared to them to be for the general 
good, and would reprobate that which appeared evil. To do 
good nnt» otliers-'-to d6 unto others, as ye would they stionld do 
unto you^is the fonndation-stone of morality. It is, therefore, 
hardly p<«Bible to exaggerate ttie importance during rude timca 


The Descent of Man. 

of the love of praise and the dread of blame. A man wlio was 
not impelled by any deep, instinctiye feeling, to sacrifico his 
!ifo for the good of others, jet was roused to such actions bj a 
sense of glory, would by his example excite the same wish for 
glory in other men, and would strengthen hy exercise the noble 
feeling of admiration. Ho might thus do far more good to Us 
tribe than by begetting offspring with a tendency to inherit his 
own high character. 

With increased esperienco and reason, man perceives the 
More remote consequences of his actions, and the self-ri^rding 
virtues, such as femperance, chastity, &e., which during early 
times are, as we have before seen, utterly disregarded, come to 
be highly esteemed or even held sacred. I need not, however, 
repeat what I have said on this head in the fourth chapter. 
Ultimately our moral sense or conscience becomes a highly 
complex sentiment— originating in the social instincts, largely 
guided by the approbation of oui fellow-men, ruled by reason, 
self-interest, and in later times by deep rehgioua feelings, and 
confirmed by instruction and habit. 

It must not be forgotten that although a h^h standard of 
morality gives but a slight or no advantage to each individual 
man and his children over the other men of the same tribe, yet 
that an increase in the niiinbor of well-endowed men and an 
advancement in the standard of morality will certainly giro an 
immense advantage to one tribe over another. A tribe including 
many members who, from possessing in a high degree the spirit 
of patriotism, fidelity, obedience, couraeo, and sympathy, were 
always ready to aid one another, and to sacrifice themselves for 
the common good, would bo victorious over most other tribes; 
and this would be natural selection. At all times throughout 
the world tribes have supplanted other tribes; and as morality 
is one important element in their success, the standard of 
morality and the number of well-endowed men wiU thiis every- 
where tend to rise and increase. 

It is, however, very difBcult to form any judgment why one 
particular tribe and not another has been euceesBful and has 
risen in the scale of civiiiaation. Many savages are in the same 
condition as when first discovered several centuries ago. As Mr. 
Bagehot has remarked, we are apt to look at prt^ress as normal 
in human society ; but history refutes this. The ancients did 
not even entei'tain the idea, nor do the Oriental nations at the 
present day. According to another high authority. Sir Henry 
Maine,' " the greatest i)art of mankind has never shewn a 

' -Anaent Law," 186], p. 32. llKhtJv Ke>:«w,' ^pril 1, 1868, p. 
For Mc. Banehot'i remarks, ' Fort 452. 


OaiP. fr'. Civilised Nations. 133 

" particle of desire tbat its civil institutions ehould be ini- 
" proTed." ProgreBS seems to depend on many concurrent 
fiiTourable conditions, far too complex to bo followed out But 
it has often been remarked, tliat a cool climate, from leading tD 
industry and to the -raiious arts, bas been highly favourable 
Hereto. The Esqnimaus, pressed by bard necesisity, have 
auccecded in many ingenious inventions, but their climate has 
been too severe for continued prepress. Nomadic habits, whether 
over wide plains, or through the dense forests of the tropics, or 
along the shores of the sea, have in every case been highly 
detrimental. Whilst observing the tmrbarous inhabitants o( 
Tierra del Fuego, it struck me that the possession of some 
property, a fixed abode, and tie union of many families under a 
chief, were the indispensable requisites for civilisation. Such 
habits almost necessitate the cultivation of the ground ; and the 
first stejffl in cultivation would jtrobably result, as I have else- 
where shevm,' from some such accident as the seeds of a fruit- 
tree falling on a heap of refuse, and producing an unasually fine 
variety. The problem, however, of the first advance of sav^^^ 
towards civilisation is at present much too difficult (o be solved, 

Natural Selection as affecting Civilised Nalions. — I have hitherto 
only considered the advancement of man from a semi-bumau 
condition to that of the modem savage. But some remarks on 
the action of natural selection on civilised nations may be worth 
adding. This subject has been ably discussed by Mr. W. E. 
Greg,'-" and previously by Mr. 'Wallacs and Mr. Galton.'" Most 
of my remarks are taken from these three authors. With 
savages, the weak in body or mind are soon eliminated; and 
those that survive commonly exhibit a vigorous state of health. 
We civilised men, on the other bond, do our utmost to check the 
process of elimination ; we build asylums for the imbecile, the 
maimed, and the sick ; we institute poor-laws ; and our medical 

' ' The Varutiod of Animals ainl 1869, and by Mr. E. Ray Lanltesier 

Flanta usdei Domestication,' vol i, in his ' Compirative Longevity,' 

p. SOB. 1870, p. 128. Similar views ap- 

' 'Fraaer'8MaEadnB,'Sept.l86a, peared previously in the -Austra- 

p.353. This article Beems to havB lasian,' July 13, 1837. 1_ har, 

rejoinder in the 'Spectator,' Oct, •• For Mr. WHllace, see ' Anttiio- 

3rd and 17th, 1868, It has also polog. EeTiew,' as before cited. Mr. 

been discussed ia the 'Q. Joumal of GaitoD in < Maoiuillan's Magazine,' 

BcJence,* 1859, p. 152, and by Mr. Ang. 1385, p. 318 ; also his greal 

Uwsoa Tail in the 'Dpblin Q, iiork, 'Hereditary Oeniai,' 1870, 
ionrual of Utdloil Sdatlce,* Feb. 

y Google 

1 34 Tlis Descent of Man. Pjbt L 

men exert their utmost sMU to save the life of every one to tlio 
lost moment. ThEie is reason to beheve that vaccination has 
preserved thousands, who from a weak eonstitutioa would 
formerly have succumbed to small-pox. Thus the weak metabera 
of civilised societies propa^te tbojr kind. No one who ha« 
attended to the breeding of domestic onimala will doubt that 
this must be highly injurious to the race of man. It is surprising 
n a want of care, or care wrongly directed, leads to the 
of a domestic race ; hut eicepting in the case of 
, hardly any one is so ignorant ss, to allow his 
worst animals to breed. 

The aid which we feel impelled to give to the helpless is mainly 
an incidental result of the instinct of sympathy, which was 
originally acquired as part of the social instincts, but sub- 
sequently rendered, in the manner previously indicated, more 
tender and more widely diffused. Nor could we check our 
sympathy, even at the ut^g of hard reason, without deteriora- 
tion in the noblest part of our nature. The surgeon may harden 
himself whilst performing an operation, for ho knows that he is 
actii^ for the good of his patient ; but if we were intentionally 
to neglect the weak and helpless, it could only be for a. con- 
tingent benefit, with an overwhelming present evil. We must 
therefore bear the undoubtedly bad effects of thewcaksurviving 
and propagating their kind; but there appears to be at least one 
check in steady action, namely that tiie weaker and inferior 
members of society do not marry so freely as the sound ; and 
this check might be indefinitely increased fay the weak in 
body or mind refraining from marriage, though tiiia is more to be 
hoped for than expected. 

In every countiy in which a large standing array is kept up, 
the finest young men are taken by the conscription or are 
enlisted. They are thus exposed to early death during war, aro 
often tempted into vice, and are prevented from marrying during 
the prime of life. On the other hand the shorter and feebler men, 
with poor constitutions, are left at home, and consequently have 
a much better chance of marrying and propagating their kind." 

Man accumulates property and l>equeaths it to his children, 
GO that the children of the rich have an advantage over the poor 
in the race for success, independently of bodily or mental su- 
periority. On the other hand, the children of parents who are 
Bhort-lived, and are therefore on an average deficient in health 
and vigour, come into their property sooner than other children, 

" Piof. H. Fick ('EiDanw der on this head, and on otker anca 
JfAtui-wisaenschail auf das R«chr,^ po;mc. 
June, 1872) haa tamo good temacka 


Cbaf, V, Civilised Nations. IJS 

and mil be likely to macrj eailier, and leave a larger ntunbei' of 
o&pnng to inheiit tbeir infeiiut couetitutions. But the m- 
heritance of property hj itself is very far &>m an evil ; for 
without the accumulation of capital the arts cculd not progress ; 
and it is chiefly through their power that the ciyilised races have 
estended, and are now everywhere est^ndioig their range, bo as 
to lake the place of the lower racea. Nor does the moderate 
accumulation of wealth interfere with the process of selection. 
When a poor man becomes moderately rich,. his children enter 
trades or .professions in which there is Bliuggle enough, so that 
the able in body and mind succeed best. The presence of a body 
of well-instructed men, who have not to labour for their daily 
bread, ie important to a degree which cannot be over-estimated; 
as all high intellectual work is carried on by tJiem, and on such 
work, material progress of all kinds mainly depends, not to 
mention other and higher advantages. No doubt wealth when 
very great tends to convert men into useless drones, but their 
number is never large; and some degree of elimination here 
occurs, for we daily see rich men, who happen tu bo fools or 
profligate, squandering away their wealth. 

Primc^eniture with entailed estates is a mora direct evil, 
though it may formerly have been a great advantage by the 
creation of a dominant class, and any goTernment is better 
than none. Most eldest sons, though thej may be weak in body 
or mind, marry, whilst the younger sons, however superior 
in these respects, do not so generally marry. Nor can worth- 
less eldest sons with entailed estates squander their wealth. 
But here, as elsewhere, the relations of civilised life are so 
complex that some compensatory checks interveno. The rcien 
who are rich through primogeniture are able to select genera^ 
tion after generation the more beautiful and charming women ; 
and these must generally be healthy in body and active in 
mind. The evil consequences, such as they may be, of tlie 
continued preservation of the same line of descent, without any 
selection, are checked by men of rank always wishing to increase 
their wealth and power; and this they effect by marrying 
heiresses. But the daughters of parents who have produced 
single children, are themselves, as Mr. Gaiton has shewn, aptto 
be sterile; and thus noble families ajo continually cut off in the 
diroct liae, and their wealth flows into some side channel ; bun 
unfortunately this channel is not determined by gupenority of 
any kind. 

Although civilisation thus checks in many ways the action of 

» 'UereditaiF Q«iuiis' 1870, pp. 132-140. 


136 The Descent of Man. Past L 

natural selection, it apparentij favours the better development 
of the body, by means of %aaK food and the freedom from occa- 
BJonal hardships. This may be inferred from civilised man 
having been found, wherever comjMired, to bo physically 
EtroDgcr than savagea." They appear also to Lave equal powers 
of endurance, as has been proved in many adventurous ex- 
peditions. Even the great luxury of the rich can be but littJf 
detrimental ; for the espeotaijoa of life of our aristocracy, at all 
ages and of both sexes, is very little inferior to that of healthy 
English lives in the lower classes." 

We will Eow look to the intellectual faculties. If in em:)i 
grade of society the members were divided into two equal 
bodies, the one including the intellectually superior and the 
other the inferior, there can be httle doubt that the fonner 
would succeed best in all occupations, and rear a greater number 
of children. Even in the lowest walks of life, skill and ability 
must be of some advantage; though in many occupations, 
owing to the great division of labour, a very small one. Hence 
in civilised nations there will be some tendency to an increase 
both in the number and in the standard of the intellectnally 
able. But I do not wish to assert that this tendency may not bo 
more than counterbalanced in other ways, as hy the multipUca- 
tioaof the reckless and improvident; but even to such as these, 
ability must be some advantage. 

It has oft«n been objected lo viewfl like the foregoing, that the 
most eminent men who have ever hved have left no offspring to 
inherit their groat intellect. Mr. Gallon says," " I regret I am 
" unable to solve the simple question whether, and Itow far, 
" men and women who are prodigies of geains are infertile. I 
'' have, however, shewn that men of eminence are by no means 
' so." Great lawgivers, the founders of beneficent religions, 
^[teat philosophers and discoverers in science, aid the progress of 
mankind in a far higher degree by their works than by leaTing 
a numerous progeny. In the case of corporeal structures, it is 
the selection of the slightly better-endowed and the elimination 
of the slightly less well-endowed individuals, and not the pre- 
servation of strongly-marked and rare anomalies, that leads to 
the advancement of a species." So it will be with the intellectual 
faculties, since the somewhat abler men in each grade of society 

" Qantrefages, ' Revne dea CoQrs 1870, p. 115. 

ScientiSquea,' 1867-68, p. 659. " 'Hereditari- Genin,,' 1870, p. 

" SetthelifthBailsiithcDluiniis, S30. 

compiled from good aothorities, in '* 'Origin of Speciea' (fifth eii- 

tbe table giren in Mr. E. E. Lan- tion, 1869), p. 104, 
kpst«r*s 'Comparative Longevity 


Chap. V. Civilised Nations. 1 37 

encceed ratber better than the less able, and consequently 
Ineroasa in ntimber, if not otherwise prevented. When in 
any nation the standard of intellect and tho mimber of intel- 
lectual men have increased, we may expect from ttie law ol 
the deriation from an average, that prodigies of gomus will, as 
ehewn by Mr. Galton, appear somewhat more frequently than 

In regard to the moral qualities, some elimication of the 
worst dispositions is always in progress even in the most oiTilised 
nations. Malefactors are executed, or imprisoned for long 
periods, so that they cannot freely transmit their bad qualities. 
Melancholic and insane persons are confined, or commit suicide. 
Violent and quarrelsome men often come to a bloody end. The 
restless who will not follow any steady occupation— and this 
relic of barbarism is a great check to eivilisation " — emigrate to 
newly-settled countries, where they prove useful pioneers. In- 
temperance is so highly destructive, that the expectation of life 
of the intemperate, at the age of thirty for instance, is only 13-8 
years ; whilst for the rural iabourers of England at the same age 
it is 40'59 years.'* Profligate women bear few children, and 
profligate men rarely marry ; both suffer from disease. In the 
breeding of domestic animals, the elimination of those individuaJa. 
though few in cumber, which are in any marked manner inferior, 
is by no means an unimportant element towards success. This 
especially holds good with injurious characters which tend to 
reappear through reversion, such as blackness in sheep; and 
with msJiiind some of the worst dispositions, which occasionally 
without any assignable cause make their appearance in families, 
may perhaps be reversions to a savage state, from which we are 
not removed by very many generations. Tliis view seems 
indeed recognised in the common espiession that such men are 
tho black sheep of the family. 

With civilised nations, as far as an advanced standard of 
morality, and an increased number of fairly good men are con- 
cerned, natural selection apparently effects but little ; though 
the fundamental social instincts were originally thus gained. 
But I have already said enough, whilst treating of the lower 
races, on the causes which lead to the advance of morality, 
namely, the approbation of our fellow-men— the strengthening 

" 'Hereditary Genius,' 1870, p. Meison's -Vital Stfltiatics.' In re. 

347. gai'd to pi'oaigacy, ees Dr. Fat/, 

'• E. Ray Laniester, 'Compara- 'InfluencB of Marriage on Hor- 

live LonEev(tJ,'5 370, p. 115. The tality,' 'Nat. Assoc for the J'romc- 

t«hle of the int-mperole a from tion of Social Science,' IS'iS. 


r38 Ttw Descent of Man. iAwi I. 

of our sympathies by habit — esample and imitation — reason — 
eKperience, and eyen self-interest — instraction during youth, and 
toligioas feelings. 

A most important obstacle in civilised countries to an uicreabH 
iu tbe number of men of a superior class has been strongly inBisted 
uu by Mr. Greg and Mr. Galton,'" namely, tie fact that the vei-y 
poor and reckless, wlio are often degiaded by vice, almost inrari- 
ably marry early, whilst the careful and frugal, who are generally 
ciherwise virtaous, marry late in life, so that they may be able 
to support themselves and their children in comfort Those who 
marry eariy produce within a, given period not only a greater 
number ot generations, but, as shewa by Dr. Duncan,* they pro- 
duce many more children. The children, moreover, that are 
bom by mothers during the prime of life are heavier and lai^er, 
and therefore probably more vigorous, than tiose txirn at other 
periods. Thus the reckless, degraded, and often vicious members 
of society, tend to increase at a quicker rate than the provident 
and generally virtuous members. Or as Mr. Greg puts the case ; 
" The careless, squalid, unaspiriag Irishman multiplies like 
" rabbits : the fingal, foreseeing, self-respecting, ambitious Scot, 
"stem in his morality, spiritual in his faith, sagacious and dis- 
" ciplined in his intelligence, paasea his best years in stru^le 
" and in celibacy, marries late, and leaves few behind him. 
"Given a land originally peopled by a thousand Saxons and a 
' thousand Celts — and in a dozen generations five-sixths of the 
" population would be Celts, bat five-sixths of the property, of 
" the power, of the intellect, would belong to the one-sixth of 
" Saxons that remained. In the eternal ' struggle for existence,' 
"it would be the inferior and iess favoured race that bad pre- 
" vailed — and prevailed by virtne not of its good qualities but of 
" its faults." 

There are, however, some checks to this downward tendency. 
We have seen that the intemperate suffer from a h^h rate of 
mortality, and the extremely profligate leave few of&pring. The 
poorest classes crowd into towns, and it has been proved by Dr. 
Stark from the statistics of ten years in Scotland," that at all 

" ' Fraaer's Magazine,' Sept. title of ' Fecnndily, Fertililj, bdiI 

1868, p. 353. 'Macmilian's Mgga- Sterililj,' 1871. See, also, Mr 

zine, Aug. 1865, p. 318. The Rev. Galton, 'Hereditary Genius," pp. 

F. W. Fanar ("Fraser'sMag.'Aug. 35L-357, foe observations to tlie 

1370, p. 264) tates a different view, above effeet. 

« ' Oo the Laws of the Fertilitr "' ' Tenth Annua) Report ol 

of WomeB,' iu 'Transact. Royal Births, Heaths, it, ^n ScotUcd, 

Soc,' Edinburgh, vol. iiiv. p. 237; 1867, p, nil. 


Chap. V. Civilised Nations. 1 39 

ages the death-rat« is higher in towns thSn in mral diatricts, 
'' \u(k during the first five, years of life the town death-rate iiS 
" lUmiXili ex^ictly double that uf the ruTjiI districta." As these re- 
tui'iiB include both the rich and the poor, no doubt more thou 
twice the number of births would be requisite to keep up the 
number of the very poor inhabitants in the towns, relatiiely to 
those in the country. With women, marriage at too early an 
age is highly injurious ; for it has been found in France that, 
" twice as many wives under twenty die in the year, as died oat 
" of the same number of the unmarried." The mortality, also, 
of bnsbands under twenty is "excessively high,"'' but what the 
cause of this may be, seems doubtful. Lastly, if the men who 
pmdentJy delay marrying until they can bring np their families 
in comfort, were to select, as they often do, women in the prime 
of life, the rats of increase in the better class would be only 
slightly lessened. 

It was established from an enormous body of stalistios, taken 
during 1853, that the unmarried men throughout France, 
between the ages of twenty and eighty, die in a much laz^r 
proportion than the married : for instance, out of every 1000 
immarried men, t>etween flie ages of twenty and thirty, 11'3 
annnally died, whilst of the married only 65 died.** A similar 
law was proved to hold good, during the years 18G3 and 1864, 
with the entire population above the age of twenty in Scotland : 
for instance, out of every 1000 uiiTnarried men, between the ages 
of twenty and thirty, W'ST annually died, whilst of the married 
only 7-24 died, that is less than half." Dr. Stark remarks on 
this, " Bachelorhood is more destructive to life than the most 
"unwholesome trades, or than residence in an unwholesome 
" house or district where there has never been the most distant 
" attempt at sanitary improvement." He considers that the 
lessened mortality is the direct result of " marriage, and the 
" more regular domestic habits which attend that state." Ho 
admits, however, that the intemperate, profligate, and criminal 
classos, whose duration of hfe is low, do not commonly marry; 
and it must likewise be admitted that men with a weak constiti\- 

" Th£3e quotalioDs are taken from tha same slrikinE paper, 

from our highest authurit}' on r,uch " I hnve taken the mean of the 

questions, namelj. Dr. Fair, In his quiuqucnaiil mean^, given in 'The 

paper 'On the Influence of Uar- Tenth Annual Report of Birthi, 

riage on the Mortality of the French Deaths, &c., in Scotiand,' 1867. 

People,' read before the Nat. Assoc. The quotation from Dr. St.itk n 

for th« Promotion cf Social Science, copied from an article in the ' Daily 

1858. Mewa,' Oct. 17th, 1868, which Dr, 

" Dr. Fan, ibid. The quota- Farr considers very carefilly wr* 


I40 The Descent of Man. Part I. 

tion, ill health, or any great infirmity in body or mind, will often 
not wish to marry, or will be reject^. Dr. Stark eeems to have 
come to the conclusion that marriage in itself is a main cause of 
prolonged life, from finding that aged married men still have a 
considerable advantage in this respect over the -unmarried of the 
same advanced age ; but every one must have known instances 
of men, who with weak healtl during youth did not marry, and 
jet have survived to old age, though remaining weak, and there- 
fore always with a lessened chance of h"fe or of marrying. There 
is another remarkahlo circumstance which seems to support Dr. 
Stark's conclusion, namely, that widows and widowers in France 
suffer in comparison with the married a very hsavy rate of mor- 
tality ; but Dr. I'arr attributes this to the poverty and evil hahita 
consequent on the disruption of the family, and to grief. On 
the wholewe may conclude with Dr. Farr that the lesser mortality 
of married than of unmarried men, which seems to be a general 
kw, "is mainly due to the constant elimination of imperfect 
■' tjpes, and to the skilful selection of the finest individuals out 
" of each Euccessiye generation ;" the selection relating only to 
the marriage state, and acting on all corporeal, intellectual, and 
moral qnalities.** We may, therefore, infer that sound and 
good men who out of pmdenco remain for a time unmarried, do 
not suffer a high rate of mortality. 

If the various checks specified in the two last paragraphs, and 
periiaps others as yet unknown, do not prevent the reckless, the 
vicious aud otherwise inferior members of society from increas- 
ing at a quicker rate than the better class of men, the nation will 
reijograde, as has too often occurred in the history of the world. 
"We must remember that progress is no invariable rule. It is 
■very difficoit to say why one civilised nation rises, becomes more 
powerful, and spreads more widely, than another; or why the 
same nation progresses more quickly at one time than at another. 
We can only say that it depends on an increase in the actual 
number of the population, on the number of the men endowed 
with high intellectual and moral Acuities, as weil as on their 
standard of esceilence. Corporeal structure appears to have 
little influence, except so far as vigour of body leads to vigour of 

It has been urged by several writers that as high intellectual 
powers are advantageous to a nation, the old Greeks, who stood 
some grades higher in intellect than any race that has ever 

■■ Dr. DnDcan remarks ('Fecund- " from the uamBiried sids to tin 

ity. Fertility,' &c., 1871, p. 334) on " marriel, leaving the unmarried 

this subject; "At everr age the " eolumm ei-owded witii (he sickly 

,'■ hcHlthy and beantifnl gu over " and unlorrtnale." 


Chap V. Civilised Kations. 141 

existed,*' ought, if the power of natnral selection were teal, to 
have tViffa still higher in the scale, increased in number, and 
stocked tte whole of Europe. Here we have the tacit assump- 
tion, so often made with respect to eorporeal structures, that 
there is some innat* tendency towards continued deTelopmont in 
mind and body. Bat developmeat of all kinds depends on many 
concurrent foTourable circumstances. Natural selection acta 
only tentatively. IndiTiduals and races may haTo acquired cer- 
tain indisputable adTsntagea, and yet have perished from failing 
in other charaeters. The Greeks may have retrc^raded from a 
want of coherence between the many small states, from the small 
size of their whole country, from the practice of slaTery, or from 
extreme sensuality ; for they did not succumb nnti] " they were 
" enervated and corrupt to the very core." ^ Tlie western nations 
of Europe, who now so immeasniably surpass their former savage 
progenitors, and stand at the summit of civilisation, owe Utile 
or none of their superiority to direct inheritance from the old 
Greeks, though they owe much to the written works of tbat 
wonderfn! people. 

Who can positively say why the Spanish nation, so dominant 
at one time, has been distanc«i in the race. The awakening of 
the nations of Europe from the dark ages is a still more perplex- 
ing problem. At that early period, as Mr. Galton has remarked, 
almost all the men of a gentle nature, ttose given to meditation 
or culture of the mind, had no refuge except in the bosom of 
a Chnich which demanded celibacy;™ and this could hardly 
fail to have had a deteriorating influence on each successive 
generation. During this same period the Holy Inquisition 
selected with extreme care the freest and boldest men in order 
to tram or imprison them. In Spain alone some of the best 
men — those who doubted and questioned, and without doubting 
there can be no prt^ess — were eliminated dnring three cen- 
turies at the rate of a thousand a year. The evil which the 
Catholic Church has thus effected is incalculable, thot^h no 
dcsbt counterbalanced to a certain, perhaps to a lai^e, extent 
in other ways; neyertheless, Europe has progressed at an un- 
paralleled rate. 

" See the ingenious and original 357) advances argomenti on the 

argnmeDt on thi> subject by Mr. other cide. SirC.Lyell had already 

Gallon, 'Hereditary Gonina,' pp. (' Principles of Geology,' vol. li, 

340-.342. _ 1868, p. 439) in a striliing passage 

Sept. 1868, p. 35J. "t the Holy Inquisition in having, 

*' 'Hereditary Genius,' 1870, fp. through eelection, lowei-ed tha gene- 

367-339. Thf Rev. F. W. Farrar ral itacdard o:" irfllignnce in £ii- 

('FraMr's Mag.; Aug. 1870, p. TOp«. 


142 Tlie Descent of Man. PiEi I. 

The lematkable success of the English as colonists, compareii 
toother European nations, has been ascribed lo their "daring 
" and persistent enei^ ; " a result whidi is well iiluetrated by 
comparing the progress of the Canadians of English and French 
extraction ; but who can say how the English gained their energy ? 
There is apparently much truth in the belief that the ■wonderful 
progiess of the United States, as well as the character of the 
people, are the results of natural selectiou ; for the more ener- 
getic, restless, and courageous men from all parts of Europe 
have emigrated during the last ten or twelve generations to tJiat 
great country, and have there sueeeeded best* Looking to the 
distant future, I do not think tb&t the Bev. Mr. Zincke takes an 
eiaggerated view when he says ;™ " AH other series of events — 
" as that which resulted in the culture of mind in Greece, and 
"that which resnlted in the empire of Rome— only appear to 
" have purpose and value when viewed in connection with, or 
"rather as subsidiary to ... . the great stream of Anglo- Sason 
" emigration to flie west." Obscure as is the problem of the 
advance of civilisation, we can at least see that a nation 
which prodnced during a lengthened period the greatest 
number of highly intellectual, energetic, brave, patriotic, and 
benevolent men, would generally prevail over less favoured 

Natural selection follows from the struggle for esisfonce ', and 
this from a rapid rate of increase. It is impossible not to regret 
Bitterly, but whether wisely is another question, the rate at 
which man tends to increase ; for this leads in barbarous tribes 
to infanticide and many other evils, and in civilised nations to 
abject poverty, celibacy, and to the late marriagesof the prudent. 
But as man suffers from the same physical evils as the lower 
animals, he has no right to expect an immimity from the evils 
consequent on the strn^le for existence. Had he not been sub- 
jected during primeval ttraes to natural selection, assuredly he 
would never have attained to his present rank. Since we see in 
many parts of the world enormous areas of the most fertile land 
capable of supporting numerous happy homes, but peopled only 
by a few wandering savages, it might be argued that the struggle 
for existence had not been sufficiently severe to force man up- 
wards to his highest standard. Judging from all that we know 
of man and the lower animals, there has always been sufBcient 
variabihty in their intellectual and moral feculties, for a steady 
advance through natural selection. No doubt such advance 

" Mr. Gaiioi, ' MaomilUn's ani! National Life,' Dec. 1 869, p. 184. 
Magazine,' August, 1865, p. 325. '" 'Last, Winter in the Dnited 

S«e Hi**, 'Nature,' 'On Darwinian! State«,' 1888, p. 29. 


Okaj. Y. Civilised Nations. 145 

■temauds maay favourable coaourrent cireumstonces ; but it may 
waU be doubted whether the most favourable wonldhayesaiBocd, 
had not the rate of tacrease been rapid, and the consequent 
struggle for existenoo eitremely severe. It even appears &om 
what we see, for instance, in parts of S. America, that a people 
which may be called civilised, saeb as the Spanish settlers, is 
liable to become indolent and to retrograde, when the con- 
ditions of life are very easy. With highly eivihsed nations con- 
tinued progress depends in a subordinate degree on natural 
selection ; for such nations do not supplant and exterminate one 
another as do savage tribes. Nevertheless the more intelligent 
members within the same community will succeed better in the 
long run tjian the inferior, and leave a more numerous progeny, 
and this is a form of natural selection. "' ~ . . 

causes of progress seem to consist of a good education during 
youth whilst the brain is impressible, and of a high standard of 
eseellence, incnlcafed by the ablest and best men, embodied in 
the laws, customs and traditions of the nation, and enforced by 
public opinion. It should, however, be borne in mind, that the 
enforcement of public opinion depends on our appreciation of 
the approbation and disapprobation of others ; and this apprecia- 
tion is founded on our sympathy, which it can hardly be doubted 
was originally developed through natural selection as one of the 
most important elements of the social instincts.^ 

On the evidence that all civUiud tiations were once barbarouB, — 
The present subject has been treated in so full and admirable a 
manner by Sir J. Lubhock,** Mr. Tylor, Mr. M'Lennan, and 
others, that I need here give only the briefest summary of their 
results. The arguments recently advanced by the Dulre of 
Argyll" and formerly by Archbishop Whately, in favour of the 
belief that man came into the world as a civilised being, and 
that all savages have since undergone degraditiou, seem to me 
w^k in comparison with those advanced on the other side. 
Many nations, no doubt, have fallen away in civilisation, and 
some may have lapsed into utter barbarism, though on this 
latter head I have met with no evidence. The Fuegians were 
probably compelled by other conquering hordes to settle in their 
inhospitable country, and they may have become in consequence 
somewhat more degraded; but it would be diflBcult to prove 

" I am niQCb indebted to Mr. " 'On the Origin of CiviliBation,' 

Jokn Morley for soma good criti- 'Proc. Ethnological Soc' Nor, 28 

ciEms on this snhject: ete, aiso, 1B67. 
Br<Ka,'LeiSdeclions,"Eeniad'AQ- " ' Prlmevil Man,' 1868. 


1^4 ^'^ Descent of Man. Pabt l. 

that they have fallen, mucli below the Botocudoe, who inhahit 
Ihe finest parts of Brazil. 

The e^ddenee that all civilised nafdons are the descendants oJ 
barharians, consists, on the one side, of clear traces of theii 
former low condition in still-exiEting customs, heliefe, language, 
i&c.; and on the other side, of proofs that savagefl are inde- 
pendently ahle to raise themselves a few steps in the scn.Io of 
ciTihsafion, and have actually thus risen. Tho evidence on the 
first head is extremely curioiis, but cannot be here given : 1 refer 
to such cases as that of the ait of enumeration, which, as Mr. 
Tylor clearly shews by reference to the words still used in some 
places, originated in counting the fingers, first of one hand and 
then of the other, and lastly of the toes. We have traces of this 
in our own decimal system, and in the Eoman numerals, where, 
after the V., which is supposed to bo au abbreviated picture of a 
human hand, we pass on to VI., &c., when the other hand no 
doubt was used. So again, " when ive speak of tliree-score and 
" ten, we are counting hy the vigesimal system, each score thus 
" ideally made, standing for 20— for ' one man ' as a Mexican or 
" Carib would put it."" According to a lai^ and increasing 
school of philoiogists, every language bears the marks of its slow 
and gradual evolution. So it is with the art of ■writing, for 
letters are rudiments of pictorial representations. It is hardly 
possible to read Mr. M'Leiman's work*" and not admit that 
almost all civilised nations etill retain traces of sueh rude habits 
as the forcible capture of wives. What ancient nation, as the 
same author asks, can be named that was originally mono- 
gamous? The primitive idea of justice, as shewn by the law of 
battle and other customs of which vestiges still remain, was 
likewise most rude. Many existing superstitions are the 
ronmants of former false religious beliefe. The highest form of 
religion — the grand idea of God hating sin and loving right- 
eousness — was unknown during primeval times. 

Turning to the other kind of evidence : Sir J. Lubbock has 
shewn that some sav^eB have recently improved a little in 
some of their simpler arts. From tho extremely curious 
account which he gives of the weapons, tools, and arts, in use 

"'Royal Institution of Great 'A Coojectnral Solntion of the 

Britain,' Maich la, 1867. Also, Origin of the Class, System o( 

' Kssearches into the Early History Relationship,' in ' Pi-do. Americ-ia 

of Mankind," ISfir,. Aiad. of Sciences,' vol, vii, Feb. 

" 'Primitive Marriage,' 1865, 1868. Prof. Kchaaffhansen ('An- 

Sea, likewise, an eicellant ai-ticle, thropolog. Review,' Oct. 1869, p 

evidently by the same author, in 373) remarks on "tha vestigea of 

the 'North British Review,' July, " haman sacrificea found both \a 

1989. Also, Mr. L. H. Morgan, " Homer and the Old Testament." 


Ghai. V. Nations. 145 

amongst savages in various parts of the world, it cajinot bo 
doubted that those have nearly all been independent discoveries, 
excepting perhaps the art of making fire.'' The Australian 
boomerang is a good instance of one such independent discovery, 
'i'he Tahitians when first visited hnil advanced in many resptcts 
beyond the inhabitants of most of the other Polynesian islands. 
There are no just grounds for the belief that the high cnltnro ol 
the native Peruvians and Mesicans was derived from abroad ;" 
maju native plants were there cultivated, and a few native 
animals domesticated. We should bear in mind that, judgiug 
from the small influence of most missionaries, a wandering creiv 
iiom some semi-civiiised land, if washed to the shores of 
America, would not have produced any marked effect on the 
natives, unless they had already become somewhat advanced. 
Looking to a very remote period in the history of the world, w<i 
Sad, to use Sir J, Lubbock's well-known terms, a paleolithic and 
neolithic period; and no one will pretend that the art of 
grinding rough flint tools was a borrowed one. In all partK of 
Europe, aa far east as Greece, in Palestine, India, Japan, Nen- 
Zealand, and Africa, including Egypt, flint tools have been 
diseovored in abundance; and of their use the existing in- 
aabitanfa retain no tradition. There is also indirect evidence of 
their former use by the Chineco and ancient Jews. Hence there 
can hardly he a doubt that the inhabitants of these countries, 
which include nearly the whole civilised world, were once 
in a barbarous condition. To believe that man ■nus abori- 
ginally civilised and then suffered utter degradation in so many 
regions, is to take a pitiably low view of human nature. It is 
apparently a truer and more cheerful view that progress na.s j 
been much mora general than retrc^ression ; that man has ri^en, j 
though by slow and interrupted steps, from a lowly condition to 1 
the highest standard as yet attained by him in knowledge, 1 
morals and religion. ' 

" Sir J. Lnbboek, ' Pi'ehistoric 

edit., 1870. 

Times,' 2nil edit. 1869, ihap. iv. 

" Dr. F. MiHle 

■od iri. rf pas^-iR. See also the 

neellent ittli c'.apter in Tybr's 

■Reise der Novu 

'Ijjtrly Hiitery if UiabiEil,' 2iid 

Theil,' Ablhoi' ii; 


The Descent of Man. 

Oil Tim AFriNiTiEB iiro Genealogy of Iiah. 

Pdltiou of mau in the animal eeries— The uatural system genesil.igicai-— 
Adaptive characters of alight Talue — Various small points of rescm- 

aj-Btem — Birthplace and antiquity of man — AbseDce of fossil coDoectiDS' 
link* — Lower atagea in the genealogy of man, as inferred, firstly fiMin 

dition of the Veitehrata— Conclusion. 

Even jf it be granted that the difforenco between man and his 
nearest allies is as great in corpet^^ stmcture as some uatu- 
raiiste mamtain, and altbongh we must grant that the dllfei^ 
ence between them is immense in mental power, yet the facts 
given in the earlier chapters appear to declare, iu the plainest 
manner, that man is descended from some lower form, notwith- 
standii^ that connecting-links have not hitherto been dis- 

Man is Uable to numerous, slight, and diversified variations, 
wbieh are induced by the same general conses, are governed 
and transmitted in accordance with the same general laws, as in 
the lower animals. Man has multiplied so rapidly, that ho lias 
cecessarily been exposed to struggle for esistenco, and con- 
seqnently to natnral selection. He has given rise to many races, 
some of which differ so much from each other, that they have 
often been ranked by naturalists as distinct species. His liody 
is constructed on the same homological plan as that of other 
mammals. He passes ttirougb the same phases of embryo- 
logical development He retains many rudimentary and useless 
structures, which no doubt were once serviceable. Characters 
occasionally mate their re-appearanco ia him, which we have 
reason to believe were poesessed by his early prc^nitors. If the 
origin of maji had been wholly different from that of all other 
animals, these various appearances would be mere empty 
deceptions ; but such an admission is incredible. These appear- 
ances, on the other hand, are intelligible, at least to a largo 
eitent, if mac is the co^iescendant with oliher mammals of soms 
unknown and lower form. 

Some naturalists, from being deeply impressed with tho 
mental and spiritual powers of man, have divided the whole 
OWBEia world into three iingdoms, the Human, tho Animal^ 


Chap. VI. Affinities and Genealogy. 147 

and tlie Vegetable, thus giving fo man a separate kingdom.' 
Spiritual powers cannot be compared or claBsed by the natu- 
ralist : but he may endeavour to shew, as I hare done, that the 
mental faculties of man and the lower animals do not di&r in 
kind, although immensely in degree. A difference in degree, 
however great, does not justify ns in placing man in a distinct 
kingdom, as will perhaps ba best illastrated by comparing the 
menfal powers of two insects, namely, a coccus or scale-insect 
and an ant, which undoubtedly belong to the same class. The 
difierence is here greater than, though of a somewhat different kind 
from, that between man and the h^host mammaL The female 
coccus, whilst young, attaches itself by its proboscis to a plant ; 
sucks the sap, but never moves again; is fertilised and lays eggs; 
and this is its whole history. On the other hand, to describe the 
habits and mental powers of worker-anfs, would require, as 
Pierre Huber has shewn, a large volume ; I may, however, briefly 
specify a few points. Anis certaioly cotmnunicats information to 
each other, and several unite for the same work, or for games of 
play. They rocoguise their fellow-ants after months of absence, 
and feel sympathy for each other. They bnild great edifices, 
keep them clean, close the doors in the evening, end post 
sentries. They make roads as well as tunnels under rivers, and 
temporary bridges over them, by clii^g together. They 
collect food for the community, and when an object, too large for 
entrance, is brooght to the nest, they enlarge the door, and 
afterwards build it up again. They store up seeds, of which 
they pz'event the germination, and which, if damp, are brought 
up fo the surface to dry. They keep aphides and other inseeta as 
milch-cows. They go out to battle in regular bands, aud freely 
sacrifice their lives for the common weal. They emigrate ac- 
cording to a preconcerted plan. They capture skves. They move 
the eggs of their aphides, as well as their own eggs and cocoons, 
info warm parts of the nest, in order that they may be quickly 
hatched; and endless similar tacts eould be given.^ On the 
whole, IJie difference in mental power between an ant and a 
coccus is immense ; yet no one has ever dreamed of pl-icing these 
insects in distinct classes, much kss in distinct king;Ioms. No 

' liidore Geoffrey St.-Hilaire gives 

of ant8 are giTen by Mr. Belt, in 

detailed account of the position 

iu his 'Naturalist in Nicaragua,' 

Bigned to man by Tarioos natural- 

1874. Sea also Mr. Moggiidge'a 

t, in their classifications: 'Hst. 

gdmirable work, 'Harvesting Ants,' 

:st. G«ii.' torn. ii. 1859, pp. liO- 

&o, 1873, also 'L'lnstjnct chei ie. 


Insectes,' by M. Geoi^e Pouehet, 

'' Some of the most interesting 

'RsYUe des Deni Mondes,' ret» 

icTs avv!- paMisbed oa the habiti 

1870 p. 682, 

t. 9 

Hosted by 


14.8 The Descent of Man. Pijii- 1. 

dowtt the difference is bridged over by other ineecls ; and this 
is not the cese ivith man and the hightir apes. But we h.ive 
overj reason to believe that the brealts in the series nro simply 
the results of many forms having become extinct. 

Professor Owen, relying chiefiy on the structure of the brain, 
has divided the mammalian series into four sub-classes. One of 
these he devotes to man; in another ho places both the 
Marsupials and the Monotremnta; so that he malies man aa 
distinct from all other mammals as are these two latter groups 
conjoined. This view has not been accepted, as far as I am 
aware, by any naturalist capable of forming an independent 
judgment, and therefore need not here be further cdnsidered. 

We can understand why a classiScation founded on any single 
character or organ— even an oi^an so wonderfully complex and 
important as tlio brain — or on the high development of Ihg 
mental faculties, is almost sure to prove unsatisfaetory. This 
principle has indeed been tried with hymonopterous insects; 
but when thus classed by their habits or instincts, the arrange- 
ment proved thoroughly artificial' Classifications may, of 
course, be based on any character whatever, as on size, colour, 
or the element inhabited ; but naturalists have long felt a 
profound conviction that there is a natural system. This 
system, it ia now generally admitted, miist bo, as &r as possible, 
genealogical in arrangement, — that is the co-descendantit of the 
aame form must be kept together in one group, apart from the 
co-descendants of any other form ; but if the parent-forms are 
related, so will be their descendants and the two groups io- 
gcthcr will form a larger group. The amount of diffircnce 
between the several groups — that is the amount of modificatioii 
which each has undergone— is expressed by such terms as 
genera, families, orders, and classes. As we hav( no record of 
the lines of descent, the pedigree can be disco\ercd oalv Ij 
observing the di^ees of resemblance betvpLon the Umgs whui 
are to be classed. For this object numerous pomts of ra^em- 
blance are of much more importance than the amount of 
similarity or dissimilarity in a few points. If two lan^agcs 
were found to resemble each other in a multitude of words and 
points of constmetion, they would be universally recognised as 
having sprung from a common source, notwithstanding that 
tbej differed greatly in some few words or points of construction. 
Bet with organic beings the points of resemblance must not 
consist oF adaptations to similar habits of Ufe ; two animals may, 
lor instance, l;avo had their whole frames modified for living in 

f VfMtToni ' I'rdttB C'ais of Insectf," vol. " IB+O, jv 87 


C'haf. YI. Affinities and Genealogy. 1<J9 

the water, and yet thoy ivill not be lironght any nearer to each 
otter in the natural Bystem. Hence we can see how it is that 
.esemhlances in several unimportant etructares, in naeleaa and 
rudimontary organs, or not now funcfionaUj active, or in au 
embryological condition, are by far the most Kerviceahle for clas- 
fiification; for they can hardly be due to adaptations within a 
late periiw ; and thus they reveal the old lines of descent or of 
true affinity. 

We can further see why a great amount of modification in 
some one character ought not to lead ns to separate widely any 
two organisms. A pait which already differs much from the 
same part in other allied forms lias already, according to tha 
theory of evolution, varied much; conseciuently it would(aslong 
as tiie organism remained exposed to the same exciting con- 
ditions) he liable to further variations of the same kind ; and 
these, if beneficial, would be preserved, and thus be continually 
augmented. In many cases the continued developmsct of a part, 
for instance, of the beak of a bird, or of the teetli of a mammal, 
would not aid tie species in gaining its food, or for any other 
object ; but with man we can see no definite limit to the con- 
tinued development of the brain and mental faculties, as fer as 
advantage ia concerned. Therefore in deteiniining tlie pc^tion 
of man in the natural or genealogical system, the eitreme de- 
velopment of bis brain ought not to outweigh a multitude of 
resemblances in other kss important or quite tmimportant 

The greater number of naturalists who have taken into con- 
sidei'ation the whole structure of man, including his mental 
faculties, have followed Blumenbach and Cuvier, and have phiced 
man in a separate Order, under the title of the Bimana, and 
therefore on an equality with the orders of the Quadrumana. 
Camivora, &c. Eecently many of oni best naturalists have 
recurred to the view first propounded by Linnieus, so remarkable 
tor his sagacity, and have placed man in the. same Order with 
Ihe Quadrumana, under the title of the Primates. The Justice of 
this conclusion will be admitted : for in the first place, we must 
Lear in mind the comparative insignificance for classification 
of the great development of the brain in man, and that the : 
stionglj-marked differences between the slnills of man and the 
Quadrumana (lately insisted upon by Bischoff, Aeby, and others" 
apparently follow from their differently developed brains. In 
the second place, we must remember that nearly all the other 
and more important differences between man and the Quadrumana 
arc manifestly adaptive in their nature, and relate chiefly to the 
eioct position of man ; such as the structure of his hand, foo^ 


1 50 TJie Descent of Man. Part I. 

and polyis, the cnivatuie of his spine, and the position of his 
head. The family of Seals offers a good illustration of the smaU 
importance of adaptive characters for classification. These 
animals differ from all other Camivora in the form of their 
bodies and in the structure of their limbs, far more than does 
man from the higher apes; yet in most systems, from that of 
CuTier to the most recent one by Mr, Flower,' seals are ranked 
as a mere family in the Order of the Camivora. If man had not 
been his own classifier, he would neyer have thought of foundit^ 
3 separate order for his own reception. 

It would bo beyond my limits, and quite beyond my knowledge, 
even to name the innumerable points of strnctnre in which man 
agrees with the other Primates. Our great anatomist and 
philosopher, Prof. Huxley, has fnlly discussed this subject,' and 
concludes that man in all parts of his organisation differs less 
from the higher apes, than these do from the lower members of 
the same group. Consec|nentIy there " ia no justification for 
" placing man in a distinct order." 

In an early part of this work I brought forward various 
facts, shewing how closely man agrees in constitution with the 
higher mammals ; and this agreement must depend on our 
close similarity in roinnte stmctoie and chemical composition. 
I gave, as instances, our liability to the same diseases, and to the 
attacks of allied parasites; our tastes in common for the same 
stimulants, and the similar effects produced by them, as well as 
by various drugs, and other such facta. 

As small unimportant points of resemblance between man and 
the Qnadmmana are not commonly noticed in systematic works, 
and as, when numerous, they clearly reveal oiir relationship, I 
will specify a few such points. The relative position of our 
features is manifestly the same ; and the various emotions are 
displayed by nearly similar movements of tie muscles and skin, 
chiefly above the eyebrows and round the mouth. Some few 
expressions are, indeed, almost the same, as in the weeping of 
certain kinds of monkeys and in the langhing noise made by 
others, during which the comers of the month are drawn back- 
wards, and the lower eyehds wrinkled. The external ears are 
curioasly alike. In man the nose is much more prominent than 
in most monkeys ; but we may trace the commencement of an 
aquiline curvature in the nose of the Hooloefc Gibbon ; and this 
in the Seiwiu/piih&Ms nasica is carried to a ridiculous extreme. 

The faces of many monkeys are ornamented with beards, 
whiskers, or moustaetes. The hair on the head grows to a greet 

' 'Pew. Zoolog. Soc.' 1863,p. 4. 

• ' Evidence as t« Man's Place in Nature,' 1863, p. '0, et jmoim. 


Ch4p. TL Affinities and Genealogy. 15I 

length in some species of SemnopithecuB ;'' and in the EoEi:et 
monkey {Maca^us radiatus) it radiateg from a point oa the crown, 
with a parting down the middle. It is commonly said that the 
forehead gives to man his noble and intellectual appearance ; bat 
the thick hair on the head of the Bonnet monkey terminates 
downwards abruptly, and is succeeded by hair so short and flna 
that at a httle distance the forehead, with the exception of the 
eyebrows, appears quite naked. It has been erroneously assertei'' 
that eyebrows are not present in any monkey. In the species 
just named llie d^ree of nakedness of the forehead differs in 
different individuals; and Eschricht states' that in oui children 
the limit between the hairy scalp and the naked forehead is 
sometimes not well defined; so that here we seem to have a 
trifling case of reversion to a pri^enitor, in whom the forehead 
had not as yet become quite naked. 

It is well known that the hair on our arms tends to converge 
from above and below to a point at the elbow. This curious 
arrangemant, so unlike that in most of the lower mammals, ja 
tommon to the gorilla, chimpanzee, orang, some species ot 
Eylobates, and even to some few American monkeys. But in 
Bylobatm agilii the hair on the fore-arm is directed downwards 
or towards the wrist in the ordinary manner ; and in B. lar it in 
nearly erect, with only a very slight f()i;ward incUnation ; so that 
in this latter species it is in a transitional state. It can hardly 
b^ doubted that with most mammals the thickness of the hair on 
the back and its direction, is adapted to throw off the rain; even 
the transverse hairs on the fore- tegs of a dog may serve for this 
end when, he is coiled up asleep. Mr. Wallace, who has carefnliy 
studied the habits of the orang, remarks that the convergence of 
the hair towards the elbow on the arms of the orang may be 
explained as serving to throw off the rain, for this animal during 
rainy weather sits with its arms bent, and with the hands clasped 
round a branch or over its head. According to lavingstone, the 
gorilla also " sits in pelting rain with his hands over his head." ' 
If the above ciplanation is correct, as seems probable, the direc- 
tion of the hair on our own arms offers a curious record of our 
former state ; for no one supposes that it is now of any use in 
throwing off the rain ; nor, in our present erect condition, is it 
properly directed for this purpose. 

It would, however, be rash to trust too much to the principle 
of adaptation in regard to the direction of the hair in man or bia 

• Uid. Geoffroy, • Hi«t. Nat. G^o.' Anat. nnd Phys.' 1837,1.51. 

l*m. ii. 1859, p. 217. • Quoted by ReBiie, ' The African 

' ' ffeber dis Eiehtnng der Sketcli Boot,' vol. i., 1873, ji. 163. 
Hanre," &c.. Mailer's 'Atohir fiiv 

y Google 

152 The Descent of Man. Part I. 

Early progenJlors ; for it is impossible to study the figures giren 
by Escbriciit of the arrai^ement of the hair on the Imman foetns 
(this being the Fame as in tlie adnlt) and not agree with this 
excellent observer tiiat other and more comples causes haro 
interrened. The points of convergence seem to stand in somo 
relation to these jwints in the embryo which are last dosed in 
during development. There appears, also, to eiist some relation 
between the arrangement of the hair oa the limbs, and the course 
of the medullary arteries.* 

It must not be supposed that the resemblances between man 
and eertajn apes in the above and many other points — such as in 
having a naked forehead, long tresses on the head, &c.— are all 
necessarily the result of unbroken inheritance from a common 
prc^enitor, or of subsequent reversion. Many of these resem- 
blances are more probably due to analogous variation, which 
ibllows,as I have elsewhere attempted to shew," from co-descended 
organisms havii^ a similar constitution, and having been acted on 
by like canses inducing similar modifications. "With respect to 
the similar direction of the hair on the fore-arms of man and 
certain monkeys, as this character is common to almost all the 
anthropomorphous apes, it may probably be attributed to in- 
heritance; but this is not eertaJn,aB some very distinct American 
monkeys are thus characterised. 

Although, as we have now seen, man has no just right to form 
a separate Order for Lis own reception, ho may perhaps claim a 
distinct Sub-order or Family. Prof. Huxley, in his last work," 
divides the Primates into three Sub-orders ; namely, the An- 
thropidte with man alone, the Simiada including monkeys of all 
kinds, and the Lemnridie with the diversified genera of lemurs. 
As far as differences in certain important points of structure are 
concerned, man may no doubt rightly claim the rank of a Sub- 
order ; and this rank is too low, it we look chiefly to his mental 
fiiculties. Nevertheless, from a genealogical point of view it 
appears that this rank is too high, and that man ought to form 
merely a Family, or possibly even only a Sub-Iamily. If we 
imagine tbree lines of descent proceeding from a common stock, 
it is quite conceivable that two of theta might after the lapse of 

• On the hair iu Hjlobates, »e Ihe Theorv of Kalnra! Sel.stion,' 

Km. Hist, of HUmmals," by C. L. 1870, p. 344. 
Martin, 1841, p. 415. Also, Isfd. •» 'Origin of Specie?,' 5th edit. 

Oeoaroyon the American monkeys 1869, p. 194. 'ITib Variation o( 

and other kinds, 'Hist. Nnt. Gin! Aaimab and Plants nndsr Domesti- 

Tol. iL 1859, p. 21B, 243. Esoh- cation,' vol. il. 1868, p. 348. 
rleht, ibid. ». 46, 55, 61. Owen, " 'Anintrodoction to the QMni- 

'Anat. of Tertebrates,' vol, ili. p. firation of Animals,' 1^89, p. 99. 
tlB. Wallace, 'ContrlbiitiDM t« 


CatP. VI, Affiniiiei mid Genealogy. 153 

Kges be so sli-ghtlj- changed as still to remain as spseies of the 
surne gernis, whilst the third line might Income so greatly 
modiliod as to deserve to tank as a distinct Sub-family, Family, 
or oven Order, But in this ease it is almost certain that the ■ 
third hne would still retain through inheritance numerous small 
points of resemblance with the other two. Here, then, would 
occur the difficulty, at present insoluble, how much weight we 
ought to assign in our classifications to strongly-marked dif- 
foreiicea in some few points, — that is, to the amomit of modifi.- 
cation undergone ; and how much to close resemblance in 
numerous unimportant points, as indicating the lines of descent 
or genoali^y. To attach much weight to the few but strong 
diSbrences is the most obTious and perhaps the safest course, 
though it appears more correct to pay great attention to the 
many small resemblances, as giving a truly natural classification. 

lu forraing a judgment on this head with reference to man, we 
rauKt glance at the classification of the Simiadss. This family is 
divided by almost all naturalists into the CatatMne group, or 
Old World monkeys, all of which are cliaract^riaed (as their 
name espresfies) by the peculiar structure of their nostrils, and by 
having four premolars in each Jaw ; and into the Platyrhine 
group or New World monkeys (including two very distinct 
sub-groups), all of which are characterised by differently 
constructed nostrils, and by haying six premolars in each jaw. 
Some other small differences might be mentioned. Now man 
unquestionably belongs in his dentition, in the structure of his 
nostrils, and some other respects, to the Catarhine or Old World 
division ; nor does he resemble the Plaljrhines more closely thau 
the Catarhines in any characters, esceptmg in a few of not much 
importance and apparently of an adaptive nature. It ia therefore 
against all probability that some New World speciessbouldhavo 
formerly varied and produced a man-lite creature, with all the 
distinctive characters proper to the Old World division; losing 
at the same time ail its own distinctive cliajractcrs. There can, 
consequently, hardly be a doubt that man is an off-shoot from the 
0!d World Simian stem ; and that under a geneaJc^cal point ot 
view, ho must be classed with the Catarhine division," 

The anthropomorphous apes, namely the gorilla, chimpanzee, 

" This is nearly the Eame classiS- adoi which anawer to theCutarbiaes. 

ration as that pi-oi'isionally adopted the Cebidai, and ths Hiipulida — 

Lv Mr. St. Geoi^B Mivarl ("TraD- these two latter groups answering 

.iet. Philosoph. Sot' 1867, p. 300), to the Platyrhinea. Mr. Mivart 

whn,8ft«r5eparatiDg the Lemnnda, «lill ahides by the same view; s^ 

dividea ths reinaiader of the Pi't- ' Natnrs,' 1871, p. 481. 
Bum into the HomiiiidiB, tb« Sini' 


: 54 ^^ Descent of Man. Past L 

onu^, and hjlobates, are b; most naturalistB separated from tlie 
other Old World monteys, as a distinct sub-gronp. I am aware 
that Gratiolet, relying on the stnictitte of the brain, does not 
' admit the existence of this sub-group, and no doubt it is a broken 
one. Thus the orang, as Mr. St. G. MiTnrt remarks," " is one of the 
" most pecuhar and aberrant forms to be found iii the Order." 
The remaining non-anthi'Opomorphous Old World monkeys, are 
again divided by some naturalistB into two or three smaller sub- 
groux>£ ; the genus Semnopitbecns, with its peculiar peculated 
stomach, being the type of one such sub-group. But it appears 
from M. Gaudrj's wonderful discoveries in Attica, that daring 
the Miocene period a form existed there, wbicU connected 
Semnopithecns and Macacus ; and this probably illustrates tbo 
manner in whicli the other and higher groups were once blended 

If tlie anthropomorphous apes be admitted to form a nntural 
fiub-group, then as man agrees with them, not only in all those 
characters which he possesses in common wiUi the whole 
Catarhine group, but in other peculiar characters, such as tha 
absence of a tail and of callosities, and ia general appearance, we 
may infer that some ancient member' of the anthropomorphous 
sub-group gave birth to man. It is not probable that, through 
the law of analogous yariation, a member of one of the other 
lower fiub^oups should have given rise to a man-liie creature, 
resembling the higher anthropomorphous apos in so many 
respects. No doubt man, in comparison with moat of his alUes, 
has undergone an extraordinary amount of modification, chiefly 
iu consequence of the great development of his brain and his 
erect position ; nevertheless, we should bear in mind that he " ia 
" but one of several esceptional forms of Primates." " 

Every naturalist, who believes in the principle of evolution, 
will grant that the two main divisions of the Simiadte, namely 
the Catarhine and Platyrhine monkeys, with their sub-groups, 
have all proceeded from some one extremely ancient progenitor. 
The early descendants of this progenitor, before they had 
diverged to any considerable extent from each other, would still 
have formed a single natural group ; but some of the epedes or 
incipient genera would have already begun to indieato by their 
diverging characters the future distinctive marks of the Catarhine 
and Platyrhine divisions. Hence the members of this supposed 
ancient group would not have been so uniform in their den- 
tition, or in the structure of their nostrils, as are the existing 


Ceas. Tl. Affinities and Genealogy. IJS 

CaSarfciae moakej^ in one way and the Platyrhiaes in another 
waj, bnt would have resemblai in this respect the allied Lemu- 
ridio, which differ greatly from each other in the form of their 
muzzles," and to an extraordinary degree in their dentition. 

The Qitarhine and Platyrhine monkeys agree in a multitude 
of characters, as is shewn by their unquestionably beloi^ng 
to one and the same Order. The many cbaractars wliich 
they possess in common can hardly have been independently 
acquired by so many distinct species; so that these characters 
most have been inherited. But a naturalist would undoubtedly 
have ranked as an ape or a monkey, an ancient form which 
possessed many characters common to the Catarhine and 
Platyrliine monkeys, otiier ctiaracters in an intermediate con- 
<lition, and some few, perhax»s, distinct from those now found in 
dither group. And as man from a genealogical point of view 
belongs to the Oatarhine or Old World stock, we must conclude, 
howeTcr much the conclusion may revolt our pride, that our 
early progenitors would have been properly thus designated." 
But we must not fall into the error of supposing that the early 
pri^enitor of the whole Simian stock, including man, was iden- 
tical with, or even closely resembled, any existing ape or monliey. 

On the Birtlipiace and Antiqtiiiy of Man. — We are naturally 
led to enquire, where was the birthplace of man at that stage of 
descent when our progenitors diverged from the Catarhino 
stock ? The feet that they belonged to this stock clearly shews 
that they inhabited the Old World ; but not Australia nor any 
oceanic island, as we may infer from the laws of geographical 
distribution. In each great region of the world the living 
mammals are closely related to the extinct species of the same 
region. It is therefore probable that Africa was formerly in. I 
habited by extinct apes closely allied to the gorilla and chim- 
panzee; and as these two species are now man's nearest allies, it 
is somewhat more probable tliat our early progenitors lived on I 
the African continent than elsewhere. But it is useless to ' 
speculate on this subject; for two or three anthropomorphous 
apes, one the Dryopithecus " of Lariet, nearly as large as a man. 

" Messrs Unrie and Mivart oa 

the Lemuroidea, 'Transact. Zaolo^. 

fichlohle,' I6SS, in which he gives 

Soc' vol. vii. 1869, p. 5. 

in detail his vienrs on tne genealcgy 

■• Haotd has come to this eame 

of man. 

tonclosioo. See 'Ueber die Ent- 

" Dr. C. Forsvth Major. ' Sor Ics 

Sinns FoBsiles tranv&i en Italie^ 


.imii.Vort.^,MS68, i. 61. Also 


rjS The Descent oj Man. Pabt I 

and closely allied to Hylobates, existed in Europe during the 
Miocene age ; and since so remote a period the earth haa 
certainly undeigone many great revolutJOES, and thtro has been 
ample lime for miprafjon on the largest scale. 

At the period and place, whenever and wherever it was, when 
man. first lost his hairy covering, he prohabiy inhabited a hot 
conntry ; a circumstance farourable for the frugiferous diet on 
which, judging from analogy, he eubsisted. %Ve are far from 
knowiiig how long ago it was when man first diverged from the 
Catarhine stock ; bat it may have occurred at an epoch as remote 
as the Eocene period; for that the higher apes had diverged 
from the lower apes as eai'Iy as the Upper Miocece period is 
shewn by the existence of the Dryojttthccus. We are also qnito 
ignorant at how rapid a rate organisms, whether high, or low in 
the scale, may be modiSed under fovorirable circumstances; wo 
know, however, that some have retained the same form during 
an enormous lapse of time. From what we see going on under 
domestication, we ieam that some of the co-descendimts of the 
eame species may he not at all, some a little, and some greatly 
changed, all within the same period. Thus it may have been 
with man, who has undergone a great amount of modification 
in certain characters in comparison with the higher apes. 

The great break in the organic chain Itetween man and his 
nearest allies, which cannot be bridged over by any extinct or 
living species, has often been advanced as a grave objection to 
the belief that man is descended from some lower form; but this 
objection will not appear of much weight to those who, from 
general reasons, behevo in the general principle of evolution. 
Breaks often occur in all parts of the series, some being wide, 
sharp and defined, others less bo in various degrees ; as between 
the orang and its nearest allies — between the Tarsius and the 
other Leniuridfe — between the elephant, and in a more striking 
manner between the Ornithorhynchus or Echidna, and aJl other 
mammals. But these breaSs depend merely on the number of 
related farms which have become extinct. At some future 
period, not very distant as measured by centuries, the civilised 
races of man will almost certainly exterminate, and replace, tho 
savage races throughout the world. At the same time tlie anthro- 
pomorphous apes, as Professor SchaafFhausen has remarked," 
will no doubt be exterminated. The break between man and Lis 
nearest aUies will then be wider, for it will intervene between 
man in a more civilised state, as we may hope, even than tlia 
Csncasian, and some ape as low as a baboon, instead of as now 
hftween the negro or Australian and the gorilla. 

■• ^nthropologistl Eevlew,' April, 1867, p. 236. 


Ghap. VI, AffirAties and Genealogy. 1 57 

With respect to the absence of iossCi remains, serving to 
lonjiect moil with his ape-like progenitors, no one will lay miicli 
stress on this fact who reads Sir C. Lyell's discussion," where 
he shews that ia oli the vertebrate clasBCS the discovery of fossil 
remains has been a very slow and fortnitous process. Nor 
shonid it be forgotten that those regions which are the raost 
likely to afford remains connecting man with some extinct apo ■ 
likq creature, have not as yet been searched by geoli^isfs. 

Lower Stages in. the Genealogy of Man, — We have seen that 
man appears to hftve diverged from the Catarhine or Old World 
division of the Simiadie, after these hnd diverged from the New ' 
World division. We will bow endeavour to follow the remote 
traces of his genealogy, trusting principally to tbe mutual 
affinities between the various classes and orders, with some 
slight reference to the periods, as far aa ascertained, of their 
successive appearance on the earth. The Lemuridie stand 
below and ceiir to the Simiadie, and constitute a very distinct 
family of the Primates, or, according io Hackel and others, a 
distinct Order. This group is diversified and broken to an 
estraordinary degree, and includes many aberrant forms. It 
has, therefore, probably suffered much extinction. Most of the 
remnants survive on islands, such as Madagascar and the 
Malayan aichipelago, where they have not been exposed to so 
severe a competition as they would have been on well-stoctod 
continents. This group likewise presents many gradations, 
leading, as Huxley remarks,** " insensibly from the crown and 
" summit of the animal creation down to creatnres &om which 
" there is but a step, as it seems, to the lowest, smallest, and 
" least intelligent of the placental mammalia." From these 
various considerafions it is probable that tbe Simiadio were 
originally developed from the progenitors of the existing 
Lemuridie ; and these in their turn from forms standiug very 
low in tbe reammallan series. 

The Marsupials stand in many important eharacters below the 
placenta! mammals. They appeared at an earlier geological 
period, and their range was formerly much more extensive 
than at present. Hence the Placentata are generally supposed 
to have lieen derived from the Implaeentata or Marsupials; 
Dot, however, from forms closely resembling the eidsting Mar- 
supials, but from their early progenitors. The Monotremata are 
plainly allied to the Marsupials, forming a third and still lower 

'Elflinenls of Geology,' 1865, '* 'Man's Place in Kaloi*, p 


15S The Descent of Man. Pakt I 

division in the great mtunmalian series. They ate represectoi 
at the present day solely by the Omithorhynehns and EcbJdca ; 
and these two terms may be safely consideied as relics of a 
much larger group, representatiTes of which have been preserved 
in Australia through some favonrablo concurrence of circum- 
stances. The Monotrematft are emiiiently interesting, as leading 
in several important points of structure towards the class of 

In attempting to trace the genealogy of the Mammalia, and 
therefore of man, lower down in the series, we become involved 
ia greater and greater obscurity; but as a most capable judge, 
Mr. Parker, has remarked, we have good reason to beiievo, that 
no true bird or reptile intervenes in the direct line of descent. 
He who wishes to see what ingenuity and knowledge can effect, 
may consult Prof Eckel's works." I will content myself with 
a few general remarks. Every evolutionist will admit that the 
five great vertebrate classes, namely, mammals, birds, reptiles, 
amphibians, and fishes, are descend^ from some one prototype ; 
for they have much in common, especially during their embryonic 
state. As the class of fishes is the most lowly oiganised, and 
appeared before the others, we may conclude that all the 
members of the vertebrate kingdom are derived from some fish- 
like anmial. The belief that animals so distinct as a monkey, 
an elephant, a humming-bird, a snake, a frog, and a fish, &c,, could 
all have sprui^ from the same parents, will appear monstrous 
to those who have not attended to the recent progress of natural 
history. For this belief imphes the former esistence of links 
binding closely together all these forms, now so utterly unlike. 

Nevertheless, it is certain that groups of animals have existed, 
or do now esist, which serve to connect several of the great 
vertebrate classes more or less closely. We have seen that the 
Ornithorhyaohua graduates towards reptiles; and Prof. Huxley 
has discovered, and is confirmed by Mr. Cope and others, that 
the Diposaurians are in many important characters intermediate 
between certain reptiles and certain birds — the birds referred 
to being the ostrich-tribe (itself evidently a widely-diffused 
remnant of a larger group) and the Areheopteryi, that strange 
Secondary bird, with a long lizard-like tail. Again, according to 

e phylum or lines of descen 

his 'Geoertlle Moiphologie * (B. ii. 
t. cliii. ond s. 435); and with more 
Mpecia! reference to man In his 

1868. Prof. Buxley, in reviewing 
this latter work ('The Academr,' 
1868, p. 42) says, that he considers 

the Veilebrata to be admirably dis- 
cussed by H^liel, although he dilfen 
on some points. He eipresses, 
also, his high estimate of the 
general tenor and spirit of th. 
whole worl:. 

Hosted by 


Chap. VL Affinities and Genealogy. 159 

Prof, Owen,™ the Ichthyosaurjans — great sea-lizards furnished 
with jjftddles — present many affinities with fishes, or rather, 
according to Hiixlej, with amphibians ; a class which, including 
in its highest divisiOTi frogs and toads, is plainly allied to tho 
Ganoid fishes. These latter fishes swarmed daring the earlier 
geological periods, and were constructed on what is called a 
generalised type, that is, they presented diyersified aflnities with 
other groups of organisms. The Lepidosiren is also so closely 
allied to amphibians and fishes, that naturalists long disputed in 
which of these two classes to rank it; it, and also some few 
Ganoid fishes, haTo been preserved from ntter estinotion by 
inhabiting rivers, whioh are harbonrs of refuge, and are related 
to the great waters of the ocean in the same way that islands 
are to continents. 

Lastly, one single member of the immense and diversified class 
of iishes, namely, the lancelet or amphioxus, is so different from 
all other fishes, that Hiickel maintains that it ought to form a 
distinct class in the vertebrate kingdom. This fish is remarkable 
for its negative characters ; it can hardly be said to possess a 
brain, vertebral column, or heart, &e. ; so that it was classed by 
the older naturalists amoi^ the worms. Many years ago Prof. 
Goodsir perceived that the lancelet presented some affinities with 
the Ascidians, which are invertebrate, hermaphrodite, marine 
creatures permanently attached to a support. They hardly 
appear like animals, and consist of a simple, toi^h, leathery 
sack, with two small projecting orifices. They belrasg to the 
Moliuscoida of Huxloy— a lower division of the great tingdom 
of theMolIusca; but they have recently been placed by some 
naturalists amongst the Yermes or worms. Their krv» some- 
what resemble tadpoles in shape," and have the power of 
swimming freely about. SLKovaleysky"* has lately observed that 
the laiTEe of Ascidians are related to the Vertebrata, in their 
. manner of development, in the relative position of the n 
system, and in possessing a stnieturo closely like the cki 
dorsaiis of verfebrata animals; and in this he has been s: 

" At the Falkland Islands 1 liad 

under s simple microscope, plainly 

tlis satisfaction of seeing, in April 

divided by tranarerse opaque parti. 

1833, and therefore some yesra be- 

tions, which I presume represent 

fore any other naturalist, the loco- 

the great cells itgul«d hy Kovalev- 

skf. At an early staeo of develop- 

dian, closply allied to Synoicnm, 

ment the tail was closely ™il5 

round the head of the lerra. 

from it. The tail was aboBt fivs 

^' 'tUmoito de I'Acad. dei 

times as long as the oblong head. 

Sciences de St. Petenhoot^,' lorn, i 

Ko. 15, isee. 

menl. it wni, s> sketched by ma 


too The Descent of Man. Pash! L 

coaflrmed jj Prof. Kupifer. M. Kovalevsky writes to me from 
Naples, that he has now carried these obserTations yet further , 
and should his results bo well eatahlished, the whole ■will form a 
(liscoverj of the very greatest value. Thus, if we may rely on 
embryology, ever the safest guide in classification, it seems that 
AVe have at last gained a clue to the source whence the Vortehrata 
were derived.^ We should then be Justified in believing 
that at an estremely remote period a group of animals esisled, 
resembling in many respects the larvaj of our present Asfiidians, 
which divei^d into two great hi-anchas— tlie one retn^rsdiiig in 
development and producing the present class of AacidiaBS,-the 
other rising to the crown and summit of the animal kingdom by 
giving birth to the Vertebrata, 

We have thus far endeavoured rudely to trace the genealogy 
of the Vertebrata by the aid of their mutual affinities. We will 
now look to man as he exists ; and we shall, 1 think, be able 
partially to restore the stnicture of our early progenitors, during 
successive periods, hut not in due order of time. This can bo 
effected by means of the rudiments which man still retains, by 
the characters which occasionally make their appearance in him 
through reversion, and by the aid of the principles of morphology 
and embryology. The various facts, to which I sliall here aliude, 
Imye-heen given in the previous chapters. 
^ The early progenitors of man must have been once covered 
with hair, both sexes liaving beards; their ears were proheblj 
pointed, and capable of movement ; and their bodies were pii)- 
vidcd with u tail, havii^ the proper mnscles. Their limbs and 
liodiea were also acted on by many muscles which now only 
occasionally reappear, but are uoiToally present in the Quadru- 
mana. At this or some earlier period, the great artery and nerve 
of the humerus ran through a supra-eondyloid foramen. The 
intestine gave forth a much larger diverticaluin or ctecum than 
that now eiisting. The foot was then prehtiiisilo, judging from 
the condition of the great toe in the fcetus ; and our progenitors, 
no doubt, were arboreal in tJioir habits, and frequented some 
warm, forest-c!ad land. The males had great canine teeth, which 

■> Rut I am boand to add 
lome competent judges dispute thb 
conclusion! f°r Instance. M. GiMd, 
In B, series of j.apers in the ' ArchiveK 

Nevenheiess, this naturalist re- 

" istence d'une cnrde doreale) chei 
" hb InverWbri par la seule con- 
"dition vltale de I'aaaptation, 
"at cette simple possibiiit^ rfn 

marks, p. 381, '• L'orgaaisadoa dj la 
"larte ascidienne en Jeh.» de 

"passage aupprmo I'-ilnmc entre 
" Iw deui sons-rtgnea, encore l.iao 

9e tout* th&irie. 


Ceap. Vj. Affinities aftd Genealogy. l6l 

Borved them ns formidable weapoiis. At a much earlier period 
the uterus was double; the escreta were voided through a cloaca; 
and the eye was protected by a third eyelid or nietifating mem- 
brane. At astiliearlieriwriod the progenitoreof man must have 
been aqTiatic in their habits; for morphology plainly telle ub that 
our lungs consist of a modiSod swim-bladder, which once serred 
<»a a float. The clefts on the neck in the embiyo of man show 
■where tho branchiffi once existed. In the lunar or weekly re- 
current periods of some of our functions we apparently still retain 
traces of our primordial birthplace, a shore washed by the tides. 
At about this same early period the true kidneys were replaced 
by the corpora wolffiana. Theheart existed as a simple pulsating 
vessel ; and the chorda dorsalis took the place of a vertebral 
column. These early ancestors of man, thus seen in the dim 
recesses of time, mnst have been as simply, or even still more 
simply organised thau the lancelet or ampMoxus. 

Tliere is one other point deserving a fuller notice. It has long 
been known that in tho vertebrate kingdom one ses bears 
rudiments of various accessory parts, appertaining to the re- 
productive system, which properly belong io the opposite sex ; 
and it has now been ascertained that at a very early embryonic 
period both sexes possess true male and female glands. Hence 
some remote progenitor of the whole vertebrate kingdom appears 
to have been hermaphrodite or androgynous." But here we 
encounter a singular difBculty. In the mammalian class the 
males possess rudiments of a uterus with the adjacent passage, 
in their vesiculs prostatieie; they bear also rudiments of 
maramffi, and some male Marsupials have traces of a marsupial 
Back." Other analogous facts could be added. Are we, then, to 
suppose that some extremely ancient mammal continued andro- 
gynous, after it had acquired the chief distinctions of its class, 
and therefore after ii had divei^d from the lower classes of the 
vertebrate kingdom ? TWs seems very improbable, for we have 
to look to fishes, the lowest of all the classes, to find any still 
existent androgynous forms.'" That various accessory parts, 

'" This is the conclusiun of pTOf. " brata are, in Ibeir early condition, 

(legenbani-, one of the highest :ih- " hcrmaphrodi'a." Similar riew* 

thoiities ia compurative anatomy; hare lonp- been hald bj- same authors, 

Ne'Crundillgederreigleich. Aa;it.' Ibongh until recently without s 

I8T0. s. 876. The result iui» Li^n firm bagif;. 

arrived at chiefly from the study of " The male Thjiaoinua offers the 

the Amphibia; but it appears from best instance. Owen, 'Anatomy oi 

the researches of Woldeycr (ai Verlebr-teB,' vol. iii. p. 771. 

quoted in 'Journal of Aoat. and " [l''rmaphrodi(lsm has been ot>- 

PKye." 1869, p. Ifll), that the seiual served in several sjwcieB of Srji raanii, 

arg-ai of ivea "the higher Tene- as well ai Id same olter fishes, 


l62 The Descent of Man. Pabt L 

propcir to each sex, are found in a rudimentary condition in the 
opposite sei, may be explained by such organs having beea 
gradually acquired by the one sei, end then transmitted in a 
more or less imperfect state to the other. When wo treat of 
^esual selection, wo shall meet with innumerable instances of 
this form of transmission, — as in the case of the spurs, plumes, 
and brilliant colours, acquired for battle or ornament by male 
birds, and inherited by the females in an imperfect or rudimentary 

The possession by male mammals of functionally impGrfect 
mammary oi^ns is, in some respects, especially curious. Tho 
Monotremata haye the proper milk-secreting glands with orifices, 
but no nipples ; and as the«^ animals stand at the very base of 
the mammalian series, it is probable that the progenitors of 
the class also had milk- secreting glands, but no nipples. This 
conclusion is supported by what is known of their manner of 
development ; for Professor Turner informs me, on the authority 
of Kolliker and Langer, that in the embryo the mammary glands 
can be distinctly traced before the nipples are in the least 
visible; and the development of successive parts in the indi- 
vidual generally represents and accords with the development of 
successive beings in the same lino of descent. The Marsupials 
differ from the Monotremata by possessing nipples; so that 
probably these organs were first acquired by the Marsupials, 
after they had diverged ftom, and risen above, the Monotremata, 
and were then transmitted to the placental mammals.™ No one 
will suppose that the Marsupials still remained androgynons, 
after they had approsimat«ly acquired their present structure. 
How then are we to account for male mammals possessing 
mammfe ? It is possible that they were first developed in the 
females and then transferred to the males; but from what 
follows this is hardly probable. 

where it is either no. 

■mal and sym- 

delle So' ' Bol g 

Dec. 28, 

lal and uni- 

1871) tht 1 ^d 


lateral. ' Dr. Zoutevi 

>eQ has given 

" Pr f G E b 

me references on this 

1 subject, more 

('Jenaiseh ^ tsch ift 

BJ tii*7- 

especially to a p^per 

hy Prof. Hal- 

212) tha tw d 

ypes of 

Wlsm^in the 'Tr 

ausaet. of the 

nipples p il h 

ghut the 

Dnteh Acad, of Soie 

several m iniii 1 

d -B. hut 

Dr. Gtmthpr doubts 

tbo fact, but 

that it q tel] (■ 

hi how both 

it has DOW been re 

corded by loo 

could h b d 

d f om the 

» to be any 

nipples f h KI p 

Is and the 

lonfsr disputed. IJ 

r. M. Lessona 

lafter f m h se f 

h Mract.*- 

writes to me, th«t 

he has Teri- 

mala. See 1 

By Dr 

tied the observatio 

Mai Hues, mam 

m rj glaadi. 

Cavollui on Serr^nus 

^ V^t EroZ 

ifaid. B. p 6 

koi hu recently eIi 

lewa ('Aocfld. 


CHir. VI. Acuities and Geitealogy. 163 

II may be suggested, as another view, that long after the 
prt^enitors of tho whole mainmaliftn class had ceased to ba 
androgynous, both sexes yielded niilfc, and thus nourished their 
young; and in tie case of the Marsupials, that both sexes carried 
their young in marsupial sacks. This will not appear altogether 
improbable, if we reflect that the males of existing sjngnathoun 
fishes receive the u^e of the females in their abdominal ponchee, 
hatch them, and afterwards, as some believe, nourish the 
young;* — that certain other male fishes hatch the e^s within 
their mouths or branchial caTitiea; — that certain male toads 
take the chaptets of eggs from the females, and wind them round 
their own thighs, keeping them there nntii the tadpoles are 
born; — that certain male birds undertake the whole duty of 
incubation, and that male pigeons, as well as the females, feed 
their nestlings with a accretion from their crops. Bat the above 
suggestion first occurred to me from the mammary glands of 
male mammals being so much more perfectly developed than 
the rudiments of the other accessory reproductive parts, which 
are found in the one sei though proper to the other. The 
mammary glands and nipples, as they esist in mala mammals, 
can indeed hardly be called rudimentary; they are merely not 
fully developed, and not functionally active. They are sympa- 
thetically affected under the influence of certain diseases, lite 
the same oi^;ans in the female. They often secrete a few drops 
of milt at birth and at puberty : this latter fact occurred in the 
curious case, before referred to, where a your^ man possessed 
two pairs of mammje. In man and some other male mammals 
these organs have been known occasionally to become so well 
developed during maturity as to yield a fair supply of milk. 
Now if We suppose that during a former prolonged period male 
mammals aided the females in nursing their offsprii^," and that 
afterwards from some cause (as from the production of a smaller 
number of young) the males ceased to give this aid, disuse of the 
organs during maturity would lead t« their becoming inactive ; 
and from two well-known principles of inheritance, this atal« of 
inaotivity would probably be transmitted to the males at the 
corresponding age of maturity. But at an earlier age these 

« Mr. Lockwood belieTeJ (ns by Prof. Wyman, in ' Proc Boston 

quoMdin'Qnart.JoHToalofSoieiice,' Sot of Nat. Hist," Sept, 15, 385T! 

April, 1S68, p. 269), from what he also Prof. Turper, ta 'Jouruia ol 

ha.1 obserrsd of the derelopment of Anat. and Phys.' Nov. I, 1S6S, p. 

Hippocnmpas, that (he vails of the 78. Dr. GUnthcr haa likewi»t de- 

ibilominal pouch of the male in scribed similar cases. 

n •' Madlle. C. Royer hne sajgetled 

r a similar view in her 'Origjie Ue 

1 rHomms • &e, 1870. 


The Descent of Man. 

twgana would be left unaflected, so that they would he almost 
equally well developed in the young of both sexes, 

Ccntdimoa. — Von Baer has defined advancement or progress in 
the oi^anic scale better than any one else, as resting on tho 
anjount of differentiajion and specialisation of the several piirta 
of a being, — when arrived at maturity, as I should be inclined to 
odd. Now aa organisms have become slowly adapted to diver- 
Bi3od lines of life by means of natural selection, their parts will 
have become more and more differentiated and speeiaLised for 
various functions, from the advantage gained by Uie division of 
physiological labonr. The same part appears often to have been 
modified first for one purpose, and then long afterwards for 
lome other and qnito distinct purpose ; and thus all the parts 
'aie rendered more and more complex. But each organism still 
retains the general type of structure of the prc^enitor from 
which it was aboriginally derived. In accordance with this 
view it seems, if wo torn to geoli^cal evidence, that organisa- 
tion on the whole baa advanced throughout tbe world by slow 
and interrupted steps. In the great kingdom of tbe Vertcbrahi 
ilihas culminated in man. It must not, however, be supposed 
that groups of organic beings are always supplanted, and dis- 
appear as soon as they have given birth to other and more 
perfect groups. The iatter, though victorious over their pre- 
decessors, may not have become better adapted for all places in 
the economy of nature. Some old forms appear to have snivived 
from inhabiting protected sites, where they have not been 
exposed to very severe competition ; and Uiese often aid us in 
constructing our genealogies, by giving us a fair idea of former 
and lost populations. But we must not fall into the error of 
looting at the existing members of any lowly-organised group as 
perfect representatives of their ancient predecessors. 

The most ancient pvi^enitors in the kingdom of the Vertebrata, 
at which we are able to obtain an obscure glance, apparently 
consisted of a group of marine animals,*' resembling the larvce of 
existing Ascidians. These animals probably gave rise to a 

" The inhibitanta of the sea- ditions for many ^.-neratiouj. eao 

ihorc must be greatly at!ected ky tardly ful to run thpir coarse iu 

the tjd«e; anlmnls living eithei- regular weekly ]>erioJs. Now It Is n 

■bout til? mean high-water inaric, mysterious fact that in the higher 

or about the mean low-watar mart, and now terrestrial Vertebrata, a* 

tuial chaa^ in a fortnight. Cou' mU and abnormal procesaea hnrc 

aeqiKntly, their food supply ivili one or mure whole weeks as their 

nndergo marked changes week \>j periods; this would oe reuderei! 

week. Tbe vital fbactions of such intelligible if the Vertebrata ai'e d.v 

tniBsls, UvtD^ uader them cos- acended bom ud alUed U 


Cnip. VI, Affinities and Genealogy. 165 

group of fishes, as lowly organised as the lancelet; and &om 
these the Ganoids, and other fishes like the Lepidosircn, must 
have been dereloped. From ench fish a Teiy small advauce 
. would carry us on to the Amphibians. We have seen that birds 
and reptiles were once intimately connected together ; and the 
Monotremata now connect mammals with reptiles In a sUght 
degree. But no one can at present say hy what line of descent 
the three higher and related classes, namely, mammals, birds, 
and reptiles, were aerived from the two lower vertebrate classes, 
namely, amphibians and fishes. In the class of mammals the 
stijps are not difficult to conceiTO which led from the ancient 
Monotremata to the ancient Marsupials ; and from these to the 
early progenitors of the placental mammals. We may thus 
ascend to the Lemuridae ; and the interval is not very wide from 
these to the Simiadie. The Simiadie then branched ofi" into two 
great stems, the New World and Old World monkeys ; and from 
the latter, at a remote period, Man, the wonder and glory of the 
Universe, proceeded. 

Thus we have given to man a pedigree of prodigious length, but 
not, it may bo said, of noble quality. The world, it has often 
been remarked, appears as if it had long been preparing for the 
advent of man : and this, in one sense is strictly true, for ha 
owes hia birth to along line of progenitors. If any single link 
in this chain had never eiistej, man would not have been exactly 
what he now is. Unless we wilfnJly close our eyes, we may, with 
our present knowledge, approximately reo^nise our parentage; 
nor need we feel ashamed of it. The most hnmble organism is 
something much higher than the inorganic dust under out feet; 
and no one with an unbiassed mind can study any living 
creature, however humble, without being struck with enthusiasm 
at its marvellous structure and properties. 

the eiistm^ tidal Ascldiaos. Many cess or function, would not, when 

instances of such periodic processes once gained, be liable to (iaugo; 

might be giveD, as the gestation of consequentlj it might be thus trans^ 

inuramals, the duration of fevers, &c. mitted through nliBost any nnmbci' 

The hatching of eggs affords also a of generations. But If the functioa 

Ci eiainple, for, according to Mr. changed, the period would have to 
tlett<'LHitd and Wat«r, Jan. 7, chauge, and would be apt to chengi. 
1871), the eggs of the pigeon are almost abruptly by a whole week, 
hatched in two weeks ; those of the This conclusion, if sound, is highly 
fonl ID three { tliose of the duck in rcmarlinble j for tha period of gesta- 
foiTj those of the goose in five; tton in each icammal, and the 
and those of the ostrich in seven hitching of each bird's eggs, and 
weeks, Ae far as we can judge, a many other vitil processes, thus 
recui'rent period, if approtimately betray lo us the primordial birth- 
el the right duration for any pro- place of these animal. 


The Descent of Mar 


On THB Races of Man. 

Tie natnrB and valne of specific characters — Application to Iha races of 
maa — Aiguuients ia fnvour of, and oppoEed to, tankiag the Eo-called 
races of msu sa distinct species — Sub-specica — Woaogeniste nnJ poly- 
genists — Convergence of character— Numei'ous points of MsemblaotB in 
body and mind between the most distinct races of man— The stale si 
man when he fimt s|>read over the earth — Each racs not descended from 
■ single paii^The ettinction of races— The formation ot rai:es— The 

. effects of crossing— Slight innuence of the dii-ect action of the con- 
ditions of life — Slight or no indnence of natural selectiou — Scsual 

It is not my intention here to describe the several so-called races 
of men ; but I am about to enquire what is the Talue of the dif- 
ferences between them under a classifioatory point of view, and 
how they liave originated. In determining whether two or more 
allied forms ought to bo ranked as spocicB or varieties, naturalists 
are practically guided by the following considerations; namely.the 
amount of difference between them, and whether such differences 
relate to few or majiy poiula of structure, and whether they are 
of physiological importance ; but more especiaOy whether tliey 
are constant. Constancy of character is what is chiefiy valued 
and sought for by naturalists. ■Whenever it can be shewn, or 
rendered probable, that the forms in question have remained 
distinct for a long period, this becomes an argument of much 
weight in fiivour of treating them as species. Even a slight 
degree of sterility between any two forms when first crossed, or 
in their ofispring, is generally considered a^ a decisive test of 
their specific distinctness; and their continued persistenca 
without blending within the same area, is usually accepted as 
sufEciect evidence, either of some degree of mutual sterility, or 
in the case of animals of some mutual repugnance to pairing. 
Independently of fusion from intercrossing, the complete 
I absence, in a well-investigated region, of varieties linking 
together any two closcly-alhed forms, is probably the most 
important of all the crlterions of their specific distinctness ; and 
this is a somewhat different consideration from mere constancy 
I of character, for two forms may be highly variable and yet not 
I yield intermediate varieties. Geographical distribution is often 
brought into play unconsciously and sometimes consciously; so 
tbat forms living in two widely separated areas, in which moet 


Ckap. VU. The Races of Mail. 167 

of ^he other inhabitants ara Bpecifically distinct, arc theuselTea 
'usually looked at as distinct ; but in truth thisafibrds no aid in dis^ 
tinguishing geographical races from so-called good or true species. 

Now let OS apply these generaUj-admitted principles to the 
races of man, viewing him in the same spirit as a naturalist would 
any other animal. In regard to tko amount of differeno© between 
the races, we must make some aJlowance for our nice powers of 
discrimination gained by the long habit of observing oui'selves. 
In India, as Elphinstone remarks, although a newly-arrived 
European cannot at first distinguish the various native races, 
yet they soon appear to him estremely dissimilar ;' and the 
Hindoo cannot at first perceive any difference between the several 
European nations. Even the most distinct races of man are 
much more lite each other in form than would at first be sup 
posed ; certain negro tribes must be excepted, whilst others, as 
Dr. itohlfs writes to me, and as I have myself seen, have 
Caucasian features. This general similarity is weE shewn by 
the French photographs in the Collection Anthropologique du 
Museum de Paris of the men belonging to various races, the 
greater number of which might pass for Enropeans, as many 
persons to whom I have shewn them have remarked. Neverthe- 
less, these men, if seen alive, would undoubtedly appear very 
distinct, so that we are clearly much influenced in our judgment 
by the mere colour of the skin and hair, by slight differences in 
the features, and by expression. 

There is, however, no doubt that the various races, when 
carefully compared and measured, differ much from each other, 
— as in the texture of thehair, the relative proportionsof all parts 
of the body,' the capacity of the lungs, the form and capacity of 
the skull, and even in the convolutions of the brain.' But it 
would be an endless task to specify the nimierous points o( 
difference. The races differ also in constitution, in aecUmatisatiou 
and in habilit; to certain diseases. Their mental characteristics 
are likewise very distinct; chiefly as it would appear in their 
emotional, but partly in their intellectual faculties. Every one 
who has had the opportunity of comparison, must have been 

' 'History of India,' 1841, vol. i. 'OntliJu;BpBcityoftlielBnga,'p.«l. 

p. 3S3. Father Eipa makes eiactly See the numercus and vat-.;Bblt 

the same remark vitb respect to table by Dr Weisbacli. from th« 

the CbiBese. observations of Dr Scherser and 

inlhropolog Theil,' 3867. 

\af. Statistics cf Amen 
Dy a A. Gould, 1869, i 



Tlie Descent of Man. 

I'ABT 1 

fitruck with tiie contrast between tho taciturn, eTcn morose, 
aborigines of S. America and tho light-hearted, talkative negroes. 
Tlicre is a nearly similar contrast between the Ulala^ and the 
Papuans,' who live under the same physical conditions, and are 
separated from each other only by a narrow space of sea. 

We will first consider the arguments which may be advanced 
in favour of classing the races of man as distinct species, and 
tLon the arguments on the other side. If a DaturaliEt, who had 
never before seen a Ni^ro, Hottentot, Australian, or Mongolian, 
were to compare them, he would at once perceive that they 
differed in a multitude of characters, some of slight and some of 
considerable importance. On enquiry he would find that they 
were adapted to live under widely different climates, and that 
they differed eomewliat in Iradily constitution and mental dis- 
position. If he were then told that hnndredsof similar specimens 
could be brought from the same conntiies, he would assuredly 
declare that they were as good speeies as many to which he had 
been in the habit of afEsing specifio names. Tliis conclusion 
would be greatly Btrengthaned ns soon as lie had ascertained that 
theso forma had all retained the same character for many 
centuries ; 8cd that negroes, apparently identical with existing 
negroes, liad lived at least 4000 years ago.* He would also hear, 
on tho autliority of an excellent observer. Dr. Lund," that the 
human skulls found in the caves of Brazil, entombed with many 
extinct mammals, belonged to the same type as that now pre- 
vailing throughout the American Continent. 

' Wnllaie, 'Thi. Malny Archi- 
pelago,' vol. ii. I8fi9, p. 178. 

' With respect to the figures in 
tha famous Egyptian cavea of Abou- 
Simbel, M. Vouchee Bays ('The 
Plurality of the Human Races,' Eng. 
-■- '"5*, p. 50), tt - ^ - 

far from 


ahle 1 

Katatlcms of tha dozen c 

man (' Races of Man.' 18S( 
speaking of 3'oiiiig Men 

formed hy Mr. Birch)) insists in 
etrongest manner that he \i iden 
in character with the Jews of 
nerp. Again, when I InokeJ al 
statue of Amunopli IIL, I ngi-eed 

=■ 20I> 

stroaEly-raarkeil r.ices 
lentilied with tW -le- 
imitv which mi trr ^a-< 



Ueasrs. Kott and UliddDii ('Types 
of Mankinil,' p. 148) state that 
KamefCi il., or the Great, has 
features superbij Bnropean; whece- 
aa Knoi, another firm heliever in 
the Bpecific distinctness of the races of 

■ As quoted by Nott and Gliddot, 
'Types of Mankind,' 1854, p. 439. 
They give also eoiToborative evi- 
dence; butC. Vo^ thinks that the 
Bab;Bcl i-eqoires further intestig*. 


Chap, VII. The Races of Man. 169 

Out naturalist would tbeu pcrliaps tnm to geograpliical dis- 
tribution, and he would probably declare that those forms must 
be distiEct species, which differ not only in appearance, bnt 
we fitted for hot, as well as damp or dry countries, and for the 
Arctic regions. He might appeal to tho fact that no species in 
the group nest to man, namely the Quadniniana, can resist a low 
temperature, or any considerable change of climate; and that 
tlie species which come nearest to man have never been reared 
to maturilj, eteu under the temperate climate of Europe. He 
would be deeply impressed with the fact, first noticed by Agassiz,' 
that the different races of man are distributed over the world in 
the same zoolt^ieal provijiees, as those inhabited by undoubtedly 
distinct species and genera of mammals. This is manifestly the 
ease with the Australian, Mongolian, and Negro races of roan ; in 
a less well-marked maimer with the Hottentots; but plainly 
with the Papuans and Malays, who are separated, as Mr. Wallace 
has shewn, by nearly the same line which divides tbe great 
Malayan and Australian zoological provinces. The Aborigines 
of America range throughout the Continent ; and this at first 
appears opposed to the above rule, for most of the productions of 
the Southern and Northern ha Ives differ widely : yet some few 
living forms, as the opossum, range from tbe one into the other, 
as did formerly some of the gigantic Edentata. The Esquimaux, 
like other Arctic animals, extend round the whole polar regions. 
It should be observed that the amoimt of difference between the 
mammals of the several zoological provinces does not correspond 
with the degree of separation between the latter; so that it can 
baldly be considered as an anomaly that the Negro differs more, 
and the American much less from the other races of man, than 
do the mammals of the African and American continents from 
the mammals of the other provinces. Man, it may be added, 
does not appear to have aboriginally inhabited any oceanic island; 
and ia this respect ho resembles the other members of his class. 

In determining whether the supposed varieties of the same 
kind of domestic animal should be ranked as such, or as spe- 
cifically distinct, that is, whether any of them are descended irom 
distinct wild species, every naturalist would lay much stress on 
the faet of their external parasites being specifically dislinct. 
All the more stress would be laid on this fact, as it would be an 
exceptional one ; for I am informed by Mr. Denny that the most 
different kinds of dogs, fowls, and pigeons, in England, are 
infested by the same species of Pediculi or lice. Now Mr. A. 
Murray has carefully examined the Pediculi coUeeted in different 

' >Dive»ity of Origlo of the ilamaa tU^e,' is the 'CtiriitLas 


170 The DescsHt of Man. PastI. 

countries from the different ra^os of man;^ ajid ha finds that 
they differ, not only in colour, but in the structure of their 
clawB and limbs. In every case in. which many specimens were 
Bbtained the differenceB were constant. The KUrgeon of a whaling 
ship in the Pacific assured me that when the Pedieuh, with 
which some Sandwich Islandera on board Bwarmed, strayed oa 
to the bodies of the English sailors, they died in the course of 
three or four days. These Pedieuh were darker coloured, and 
appeared different from those proper to tbe natives of Cbiloo in 
South Ametii^, of which he gave me specimens. These, again, 
appeai'cd latter and much softer than European lice. Mr. 
llnrray procured foni kinds from Africa, namely from the Negroes 
of the Eastern and Western coastB, from the Hottentots and 
Kaffirs; two kinds from the nativeeof Australia; twofromNorth 
and two from South AmericBt In these latter cases it may be 
presumed that the Pedicnli came from natives inhabiting different 
districts. With insects slight structural differences, if constant, 

. are generally esteemed of siwcific value : and the fact ot the 
races of man being infested by parawtea, which appear to bo 
specifically distinct, might fairly be urged as an argument that 

] the races themselves ought I0 be classed as distinct species. 

Onr supposed naturalist having proceeded thus far in his 
investigation, would next enquire whether the races of men, when 
crossed, were in any degree sterile. He might consult the work' 
of Professor Broca, a cautions and philosophical observer, and in 
this he would find good evidence that some races were quite 
fertile t<^ther, but evidence of an opposite nature in regard to 
other races. Thus it has been asserted that the native women of 
Australia and Tasmania rarely produce children to European 
men ; the evidence, however, on this head has now been shewn 
to be almost valueless. The half-castes are killed by the pure 
blacks : and on account has lately been published of eleven half- 
caste youths murdered and burnt at the same time, whose 
remains were found by the polica'" Again, it has often been 
said that when mulattooa intermarry they produce few children ; 
on the other liand. Dr. Bachman of Charleston" positively 

I. A. 

('Revne des Cours Scientifiques,' 
March 1366, p. 239) much eridence 
Mr. T. A. Murray, in the * Anthi-o- that Australians and Europeans ar« 
• )g. Review," April 1868, p. liii, not sterile when crossed. 
.... ,...._ ^._„ o. — ...I.-.. 11 >^jj E^ncnination of ProE 

AgaEsii's Sketch of the Nat. Pro- 

■ransact. R Soc. of Ediahui^h,' 

who hare borne children to a 

■ii. 1861, p. 56T. 

hi the Phenomena of Hybridity 

their own race, is disproved. 

Genna Homo,' Eng. translal. 


- Hostedby VjOOQIC 

CJhap. Vn. The Races if Man. 171 

assorts that he has known mulatto famihee which havo inter- 
married for aereral generations, and have continued on an 
average as fertile as either pui'o wliites or pure blacks. Enquiriea 
formerly made by Sir 0. Lyell on this subject led him, as ha 
informs me, to the same conclusion." In the United States tha 
census for the year 1854 included, according to Dr. Bachman, '" 
405,761 mulattoes ; and this number, considering all the circum- 
stances of the case, seems small ; but it may partly be accounted 
for by the degraded and anomalous position of the class, and by 
the profligacy of the women. A certain amount of absorption of 
mulattoes into negroes must always be in progress; and this 
would lead to an apparent diminution of the former. The inferior 
Titality of mulattoes is spoken of in a trustworthy work'* as a 
well-known phenomenon ; and this, although a different considera- 
tion from their lessened fertihty, may peihaps be advanced as 
a proof of the specific distinctness of the parent races. No donbt 
both animal and vegetable hybrids, when produced from extremely 
distinct species, are UaWe to premature death; but the parents 
of mulattoes cannot be put under the category of extremely 
distinct species. The common Mule, so notorious for long life 
and vigour, and yet so sterile, shews how little necessary con- 
nection there is in hybrids between lessened fertility and vitality ; 
other anaJogous eases could be cited. 

Even if it should hereafter bo prored that all the races of 1 
men were perfectly fertile together, he who was inclined from ! 
other reasons to rank them as distinct species,might with justice | 
argue that fertility and steribty are not safe criterions of specific ' 
distinctness. We know that these qualities are easily affected 
by chained eonditions of life, or by close inter-breeding, and that 
they are governed by highly complex laws, for instance, that of 
the unequal fertility of converse crosses between the same two 
species. With forms which must be ranked as undoubted 
species, a perfect series esists from those which are absolutely 
sterile when crossed, to those which are almost or completely 


TiDcesof theADimal Woi-Id,' Charles- 

the children are few and sioKl 

ton, 185j, p. 44. 

This belief, as Mr. Reads romari 

" Dr. Kobife writes to ma thit 

deserves atUntJan, a» white m 

he found the miied races in tho 

hare visited and resided on the Gt 

Great Sahara, derived from Arabs, 

Coast for fonr hundred years. 

BerbeiB, an* Negroes of three tribes, 

eiiraordiaarily feitile. On the other 

time t« gain knowledge throu 

haad, Mr. ffinwood Reads infurms 


me that the Negroes on the Gold 

" ' Military and Anthropo^ 

Coast, though admiring white men 

Statistics of American Soldiers.' 

and mulattoes, h.ive a maiim that 

B. A. Gould, 1869. p. 319. 

lonlattoeB should not intermarrj, •» 



The Descent of Man. 


fertile. Tho dt^rees of sterility do not coincide strictly with 
the degrees of difference between tho parents in eiterual structure 
Br habits of life, Man in many respects may be compared with 
those animals which have long been domesticated, and n largo 
body of evidence can be advanced in favour of the Pallasisn 
doctrine," that domestication tends to eliminate the sterility 
which is so general a result of the crossing of species in a state 
of nature. From these several considerations, it may be justly 
I ui^d that the perfect fertility of the intercrosBed races of man, 
lif established, would not absolutely preclude ns from ranking 
I them as distinct species. 

Iniiependentlj- of fertjhty, the characters preseuttd by the off 
spring from a cross have been thongtt to indjcatt whether or not 
the parent-forms ought to be ranked as species or varieties, but 
after carefully studying tho evidence, I have come to the con 
elusion that no general rules of this kind can be tru'.ted The 

ordinary result of a cross is the production of a blended or 

I* 'The Vari.-itioii of Animals nnd 

sterile, it IS scarcely possible that 

Plants under DomestieatidD ' vol ii 

their sterility should be augmenleo 

p. 109. I msy here remind be 

by h preseivaton or sarvlval of 

reader that the sterility of pec ea 

h m nnd more sten'e indi- 

when crossed Is Dot a p 

dnals for as the sfeiihty in- 

acquired qualHj\ but, lik h 

<xs, fewer and fewer offapnng 

b produced from nhich to 

ed together, is incidental n h 

b d, and at last only single m 

d Tidu will be produced, at the 

of Ibese diiferenCBS is unkn wn b 

n ntervoU But there is eiea 

Ihey relate moiB especially to the re 

a higher giade of Eteniitr thau 

productive tystem and much less so 

this Both Gartner and kolrenter 

to eiternai structure ir to ordinary 

hive proved that lU genera of plants 

differences in conslitition One 

inclndug many species a ser » 

impoitant element n the sterility 

cm be lormed from species which 

(,( crossed ii ecie^ apgi lentlv lies in 

when crossed yieli lewer and fewer 

one or bcth i ay ng been long babi 

seeds to spec es n hich uevei p o 

mated to filed eondit on> tor we 

duCB a single seed but yet are 

IcDow that changed conditions have 

aflectcd by the pollen of the other 

a special mfluence on the repro 

spec es as shewn by the swell ng 

ductiTC system and we have good 

o. the geimen It i. here manl 

easnn to believe (as beEnre re 

tctlj impossible to select the mo e 

marked) that the liu<.tuntlng cod 

stenle individuals which have ai 

dltioDa of domfttication lend to 

ready censeil to yirl 1 seeds , so that 

'jliminate tbal'stenWy which is «> 

the acme of sterilitj when the 

general with species in a mlural 

gennea alone is aftecleJ cannot 

stale when crosseJ It ha< else 

have beeu gained throggh selecti b. 

where been shewn by me (ibid voL 

This acme, and no doubt the other 

11 p 18j and Ongm of =lpecies* 

gradesofstenlity re the mciJentsJ 

5.J1 edit. p. ai7), that the .terility 

resulU of certain unknown differ- 

Hosted by GoOC^Ie 

Chap. Vn. The Races of Man. 1 73 

intermediate form ; but in certain cases some of the oflfepring take 
closelj after one parent-form, and some aft«r the other. Tiiis is 
OBpecially apt to occur when the parents differ in characters 
lehich first appeared as sudden Yariations or monBtrosities." I 
refer to this point, becanse Dr. Eolilfs infoi-ms me that be hae 
frequently seen in Africa the offspring of negroes crossed with 
members of other races, either completely black or completely 
white, or rarely piebald. On the other hand, it is notorioua 
that in America mnlattoes commonly present an intermediate 

Wo have now seen that a naturalist might feel himself fully 
justified in ranking the races of man as distinct species ; for ho 
has found that they are distmguished by many differences in 
structure and constitution, some being of importance. These 
differences have, also, remained nearly constant for very long 
periods of time. Our natnialiat will have been in some degree 
influenced by the enormous range of man, which is a gi'eat 
anomaly in the class of mamma,la, if mankind be viewed as a 
sii^la species. He will have been struck with the distribution of 
the several so-called races, which accords with that of other 
undoubtedly distmet species of mammals. Finally, he might 
m^e that the mutual fertility of all the races has not as yet been 
fully proved, and even if proved would not be an absolute proof 
of their specific identity. 

On the other side of the question, if our supposed naturalist 
were to euqiiire whether the forms of man keep distinct like 
ordinary species, when mingled together in large numbers in the 
same country, he would immediately discover that this was by 
no means the case. In Brazil he would behold an immense 
mongrel population of Negroes and Portuguese ; in Chiloe, and 
other parts of South America, he would behold the whole popu- 
lation consisting of Indians and Spaniards blended in various 
degrees." In many parts of the same continent he would meet 
with the most complex crosses between Negroes, Indians, and 
EuropEjans ; and judging from the vegetable kingdom, such triple 
crosses afford the severest test of the mutual fertility of the 
pa/ent-forms. In one island of the Paciiic ho would find a 
small population of mingled Polynesian and English blood ; and 
in the Eiji Archipelago a population of Polynesian and Negritos 

" 'Thf VariaUon of Auiraab,' bucmss and energy of tlio PaulistM 

" Jl' de'Qiwtref^es has p'*" race of Fortugues* and Indians, witli 
(' Anthropolog. Keritw,' Jan. 1869, a miitui-e of the blooii of othei 
p. Si) >D interesting account of the ibobl 


174 The Descent of Man. PitiT 1. 

orosGed in all degrees. Many analogoos cases could be be adaed ; 
for instancB, in Africa. Hence the races of man are not suf- 

i ficiently distinct to inhabit llie same country without fusion ; 

land the absence of fnsioQ affords the 1:81181 and beat test oi 

japDcific distinctness. 

Our naturalist would likewise be much disturbed as soon aa 
ho perceived that the distinctive characters of all the races were 
highly variable. This fact strikes erery one on first beholding 
the negro skves in BroBil, who have been imported from :ill 
parts of Africa. The same remark holds good with tlie 
Polynesians, and with many other races. It may be doubted 
whether any character can be named which is distinctive of a 
race and is constant. Savages, even within the limits of the 
same tribe, are not nearly so uniform in character, as lias l)een 
often asserted. Hottentot women offer certain peculiarities, 
more strongly marlied than those occurring in any other race, 
but these are known not to be of constant occurrence. In the 
several American tribes, colour and hairiness differ considerably; 
as does colour to a certain degree, and the shape of the features 
greatly, in the Negroes of Africa. The shape of the skull varies 
much in some races;" and so it is with every otier character. 
Now all naturalists have learnt by dearly-boaght expuience, how 
rash it is to attempt to define species by the aid of inconstant 
But the most weigbty of all the ailments against treating 

I the races of man as distinct species, is that they graduate into 

leach other, independently in many cases, as far as we can judge, 
'of their having intercrossed. Man has been studied more 
oarefally than any other animal, and yet there is the greatest 
possible diversity amongst capable judges whether he should be 
classed as a single species or race, or as two (Tirey), as three 
(Jacqninot), as four (Kant), five (Blumenbach), six (Buffon), 
seven (Hunter), eight (Agassiz), eleven (Pickering), fifteen 
(Bory St. Vincent), sixteen (Desmonlins), twenty-two (Morton), 
sisty (Crawfurd), or as sixty-three, according to Burke.'' This 
diversify of jui^ment does not prove that the races ought not 
to be ranked as species, but it shews that they graduate into each 

giues of America ind Anatralii. subject in Wait(, 'Inlroduet, M 

Prtf. Huxlev mjn (' Transact. Inter- Anthropology,' Eog. transit. 186:t, 

nat. Uongies* of fnbUt. Arch.' pp. 198-208, 227. J haTe takpp 

1868, p. 105) that the sknlls of some of the above statelDtnts from 

inaDir Sooth Cermans and Swiss are H. Tuttle'g •Origin and Aatiquilf 

" as short and aa hroad as these of of Phj«cal Man,' Boslon, 1866, p, 

" tht I'artars," Ac. 35. 


Ch*p. VH, The Races of Man. i7S 

otiier, and that it is hardly possible to discover clear distinctive \ 
chanuiters between them. 

Every naturalist who has had the misfortune to undertake the 
description of a gionp of highly varying organiEms, has en- 
couEterod cases (I speak after experience) precisely like that of 
man ; and if of a cautions disposition, he will end by uniting all 
the forms which graduate into each other, nnder a single 
species ; for he will say to himself that he has no right to give 
names to objects which he cannot define. Cases of this kind 
oocnr in the Order which includes man, namely in certain genera 
of monkeys; whilst in other genera, as in Cereopithecus, most of 
the species can be determined with certainty. In the American 
genus Cebns, the vorions forms are ranked by some naturalieta 
as species, by others as mere gec^raphical races. Kow if 
numerons sx)ecimena of Cebns were collected from all parts of 
South America, and those forms which at present appear to be 
specifically distinct, were found to graduate into each other by 
close steps, they would usually be ranked as mere varieties or \ 
races; and this course has been followed by most naturalists 
with respect to the races of man. Nevertheless, it mnst be 
confessed that there are forms, at least in the vegetable king- 
dom," which we cannot avoid namii^ as species, but which are 
connected together by numberless gradations, independently of 

Some natnialists have lately employed the term " snb-specios" 
to designate forms which possess many of the characteristics of 
true species, but which hajrdly deserve so high a rank. Now if 
we reflect on the weighty arguments above given, for raising tlie 
races of man to the dignity of species, and the insuperable difB- 
ciilties on the other side in defining them, itseems that the term 
" snb-speeies " might here be used with propriety. But fromj 
long habit the term " race " will perhaps always be employed.! 
The choice of terms is only so far important in that it is desirabloj 
to use, as fer as possible, the same terms for the same di^rees of' 
difference. Unfortunately this can. rarely be done : for the larger 
genera generally include closely-allied forms, which can be 
distinguished only with mneb difflcnlty, whilst the smaller 
genera within the same family include forms that are i)erfectly 
distinct ; yet all mnst be ranked equally as species. So again, 
species within the same large genna by no means resemble 
each other to the same degree : on the contrary, some of them 

■• Piof. Fdgfli 

has oarefnlly Je- 

mribed uveral st 

rikins cuses ia his 

'BotBBlBdia Mitt 

heiluDgen,' B. ii. 

leeS, 1. 294-369. 

VtoL Asa Gray 


176 The Descent of Man. Pjbt 1 

can generally be arranged in little groups round other species, 
like satellites round planeta.'" 

The question whether niiiiikind eoDKista of one or severa 
species has of late jcarsbeen muclidiscussed by anthropologists, 
who are diTJded into the two schools of mouogenists and 
polygenists. Those who do not admit the principle of evolution, 
must look at species as separate creations, or as in sorce manner 
as distinct entities; and they must decide what forms of man they 
will consider as species hy the analogy of the method commonly 
puiBUGd jn ranking other organic beings as species. Bnt it is a 
hopeless endeavour to decide this point, until some definition of 
the term "species" is generally accepted; and the definition 
must not include an indeterminate element such as an act of 
I creation. We might as well attempt without any definition to 
decide whether a certain number of houses should be called a 
viEage, town, or city. We have a practical illustration of the 
difficulty in the nevei'-ending doubts whether many closely-allied 
mammals, birds, insects, and plants, which represent each 
other respectively in North America and Europe, should bo 
ranked as siwcies or geographical races ; and the Hko holds true 
of the productions of many islandssituated at some little distanco 
from the nearest coutinent. 

Those naturalists, on the other baud, who admit the prindpit 
of evolution, and this is now admitted by the majority of risinc 
men, will feel no doubt that all the races of man are descended 

Ifrom a single primitive stock; whether or not they may tliiuk 
fit to des^inate the races as distinct species, for the sake of ex- 
pressing their amount of difference,^' With our domestic 
animals the question whether the various races Lave arisen from 
one or more species is somewhat differenL Although it may be 
admitted that all the races, as well as all the na^ral species 
within the same genus, have sprung from the same primitive 
stock, yet it is a fit subject for discussion, whether all the 
domestic races of the dog, for instance, have acquired thcii 
present amount of difference since some one species was firjl 
domesticated by man ; or whether they owe some of thiir 
characters to inheritance from distinct species, which had 
already been differentiated in a s(at« of nature. With man no 
sneh question can. arise, for he cannot be said to have beta 
domesticated at any particular period. 
During an eariy stage in the divergence of tlio racus of man 

" 'Orisin of Swcies,' 5th f4it. in tha 'Fortuigbtly lieii-!«,' I8S5, 
^ 68. p. 375. 

" Sot F.-oC Huiky to tbiii cSkI 


Ohap, Vll. The Races of Man. IJJ 

from a eommon stock, the differences between the races ami 
thair number miist have been Email ; conseqnentlj oa far as 
their distingimiing characters are concerned, they then had Icks 
claim to rank as distinct species than the existing so-cai led races. 
NeTortholess, so arbitrary is the term of species, that snch early 
races would perhaps havo been ranked by some natnralista as 
distinct species, if their differences, although extremely slight, 
had been more constant than they are at present, and had not 
graduated into each other. 

It is however possible, thot^h far from probable, that the 
early progenitors of man might formerly have diverged mnch in 
character, until they became more unlike each other than any 
now existing races; but that subsequently, as suggested by 
Vogt,'" they convolved in character. When man selects the off- 
spring of two distinct species for the same object, he sometimes 
induces a considerable amount of convergence, as far as general 
appearance is concei'ned. This is the case, as shewn by Von 
Nathusius,™ with the improved breeds of the pig, which are 
descended from two distinct species; and in a less marked 
manner with the improved breeds of cattle. A great anatomist, 
Gratiolet, maiatains that the anthropomorphous apes do not 
form a natural sub-gronp ; but that the orang is a highly 
developed gibbon or semnopitheous, the chimpanzee a highly 
develoiKjd maca«ua, and the gorilla a highly developed mandrill. 
If this conclusion, which rests almost esolusively on brain- 
cliaractors, be admitted, we shonld have a case of convei^enco 
at least in estemal characters, for the anthropomorphous ai)cs 
are certainly more like each other in many points, tlian they are 
to other apes. All analogical resemblances, as of a whale to a 
fish, may indeed be said to be cases of convergence; but this 
term has never been applied to superficial and adaptive resem- 
hlanccs. It would, however, be extremely rash to attribute to 
convei^nce close similarity of character in many points of 
structure amongst the modified descendants of widely distinct 
beings. The form of a crystal is determined solely by the molo- 
cular forces, and it is not surprising that dissimilar substances 
shonld sometimes assume the same form ; but with oi^»nic 
beings we shonld bear in mind that the form of each depends on 
an infinity of comples relations, namely on variations, due to 
causes far too intricate to be followed,— on the nature of the 
variations preserved, these depending oa the physicial condi- 

« ' Lectures on Man," Eng, fr 
hit. I8S4, p. 4eS. 

II iT Hosted by Google 

i;8 The Descent of Man. Paki- 1. 

tions, and still more on lie surroimiJ^cg organisms wljioli com- 
pete with each,— and lastly, on inheiitanceCinitseif a fluctuating 
element) from innnmerable progenitors, all of ivbich have had 
their forma determined throuRh equally comploi relations. It 
appears incredible that the modified descendants of two oipin- 
isnis, if these difiered from ea*h other in a marked manner, 
shoald erer aft«rwarda conTorge so closelj as to lead to a near 
approach to identity throughout their whole organisation. In 
the case of the convergent races of pigs above referred to, evi- 
dence of their descent from two primitive stocks is, according to 
Von Nathnsius, still plainly retained, in certain bones of their 
skulls. If the races of man had descended, as is supposed by 
some naturalists, from two or more species, which differed from 
each other as much, or nearly as much, as docs the orang from 
the gorilla, it can hardly bo doubted that marked differences in 
the structui'e of certain bones would still be discoverable in maji 
as ho now osists. 

, Although the existing races of man differ in many respects, as 
/in colour, hair, shape of skull, praportions of the body, (fee, yet 
I if their whole structure be taken into consideration they are 
' found to resemble each other closely in a multitude of points. 
Many of these ace of so unimportant or of so singular a nature, 
that it is extremely improbable that they should have been inde- 
pendently acquired by aboriginally distinct species or races. 
The same remark holds good with equal or greater force with 
respect to the numerous points of mental similarity between the 
most distinct races of man. The American aborigines, Negroes 
and Europeans ace as different from each other in mind as any 
three races that can be named; yet I was incessantly struck, 
whilst living with the Fuegians on board the " Beagle," with the 
many little traits of character, shewing how similar their minds 
■were to ours; and so it was with a full-blooded n^ro with whom 
I happened once to be intimate. 

He who will read Mr. Tyler's and Sir J. Lubbock's interesting 
works " can hardly fail to be deeply impressed with the dose 
similacity between the men of all caces in tastes, dispositions and 
habits. This is shewn by the pleasuce which they aU take in 
dancing, rude music, acting, painting, tattooing, and otherwise 
decorating themselves ; in their mutual comprehension of gesture- 
language, by the same espression in their features, and by the 
same inartieulale cries, when excited by the same emotions. 
This similarity, or rather identity, is striking, when contrasted 


Chap. VII. The Races of Man. 179 

with tho different cspressiona and cries made bj distinct Bpecies 
of monkeys. There is good evidence that the art of shooting 
with bows and arrows has not been handed down from any 
common progenitor of mankind, jet as Westropp and Nilsson 
have remarked/" the stone ariow-heads, brought from the most 
distant parts of the world, and raaund'actnred at tho most remote 
periods, are aJmost identical ; and this fact can op 'y I" fi-ei^-^T.\pA \ 
for h y the various racea ha^g siniilBr taTentiT " ftr mpnta.l ! ^ 
po wMS . Tho same observation has been made by arehseoloeists ^ 
wKffrespect to certain widely-prevalent ornaments, such as zig- 
zags, &C. ; and with respect to various simple hehefs and cus- 
toms, such as the burying of the dead under megalithic struo- 
turea. I remember olworving in South America," that there, as 
in so many other parts of the world, men have generally chosen 
the summits of lofty hills, to throw op piles of stones, either as 
a record of some remarkable event, or for burying their dead. 

Now when naturalists olraerre a close agreement in numerous 
small details of habits, tastes, and dispositions between two or 
more domestic races, or between Loarly-aUied natural forms, 
they use this tact as an argument that they are descended from a 
common progenitor who was thus endowed; and consequently 
that all should be classed under the same species. The same 
argument may be applied with much force to the races of man. 

As it is improbable that the numerous and unimportant points 
of resemblance between the several races of man in bodily struc- 
ture and mental faculties (I do not here refer to similar customs) 
should all have been independently acquired, they must Lave been 
inherited from progenitors who had these same characters. We 
thus gain some insight into the early state of man, befo^re he had 1 
spread step by step over the face of the earth. The spreading n 
of man to regions widely separated by the sea, no doubt, pre- 1\ 
ceded.any great amount of divergence of character in the several 1 1 
races; for otherwise we ahonld smnetimea meet with the same \ I 
race in distinct continents ; and this is never tho case. Sir J. I 
Lubbock, after comparing the arts now practised by savages in 
all parts of the world, specifies those which man could not have 
known, when he first wandered from his original birth-place; 
for if once leamt they would never have been forgotten.^ He 

I' 'On AanlDgous Forma oi Im- ' jDnrual of EtbnolDgical Sac' u 
plements.' io 'Memoirsof ADthropo- given in 'BcrentiSc Opinion,' Jnne 
log. Soc.' by H. M. Westropp. 'Ths 2od, t869, p. 3. 

navia,' Eng- translat. ediWd by Sir o(" tbe " Be^le," ' p. 46. 
4. Lubbock, 1868, p. IM. " 'Frehistotie Times,' 1869, p, 

<• Westropp, ' On Cromlechs,' &«., 574. 


l8o Tli£ Descent of Man. PartL 

tlina shews that " the spear, which is but a dovelopffient of the 
•' knife-point, and the club, which is but a long hajumer, aro the 
" only tiling left," He admits, however, that the art of making 
fire probably had been already discovered, for it is common toi 
all the races now existing, and was known to the ancient cave- 
inhabitants of Europe. Perhaps the art of making nide canoes 
or rafts was likewise known ; but as man existed at a remote 
epoch, when the land in many places stood at a very different 
level to what it does now, he would iave been able, without the 
aid of canoes, to have spread widely. Sir J, Lubbock further 
\ remarks how improbable it is that our eai'liest ancestors could 
\ have " counted as high as ten, considering that so many races 
Vnow in existence cannot get beyond four." Nevertheless, at 
this early period, the intellectual and social faculties of man 
could hardly have been inferior in any extreme degree to those 
: possessed at present by the lowest savages ; otherwise primeval 
,' man could not have been so eminently successful in the struggle 
for life, as proved by his early and. wide diffusion. 

From the fundamental differences between certain languages, 
some philologists have interred that when man first became 
widely diffused, he was not a speaking animal; but it may be 
suspected that languages, far lesB perfect than any now spoken, 
aided by gestures, might have been used, and yet have left no 
traces onsubseqnentandmore highly-developed tongues. With- 
out the use of some language, however imperfect, it appears 
doubtful whether man's intellect could have risen to the 
standard implied by his dominant position at an early period. 

Whether primeval man, when he possessed but few arts, and 
those of the rndest kind, and when his power of language was 
extremely imperfect, would have deserved to be called man,must 
depend on the definition which we employ. In a series of forms 

|>{raduating insensibly from some ape-hke creature to man as he 
now exists, it would bo impossible to fix on any definite point when 
the term " man " ought to be used. But this is a matter of very 
little importance. Bo again, it is almost a matter of indifference 
whether the so-called races of man are thus designated, or are 
ranked as species or sub-species ; but the latter term appears the 
more appropriate, finally, we may conclude that when the 
principle of evolution is generally accepted, as it surely will be 
before long, the dispute between the monogenists and the poly- 
genists will die a silent and unobserved death. 

One other question ought not to be passed over without notice, 
namely, whether, as is sometimes assumed, eaeh sub-species or 
race of man has sprung from a single pair of progenifors. With 


Chap. VII. The ExtinCiLn of RacsS. iSl 

our domestic onimals a new race can readily be formed by care- 
fully matcLing tiie varying offspring from a siEglo pair, or eren 
from a single individual possessing some now character; but 
most of our races have been formed, not intentionally from a ] 
selected pair, but unconsciously by the preservation of many in- 1 
dividualawidclihavevarjed, however slightly, in some nsoful or | 
desired manner. If in one country stronger and heavier horses, 
and in another country ligjitor and fleeter ones, were habituiUy 
preferred, wa may feel sure tJiat two distinct sub-breeds would 
be produced in the course of time, without any one pair having 
been separated and bred ivo\n, in either country. Manyraceai 
have been thns formed, and their manner of formation is closely ' 
analogous to that of natural species. We know, also, that the 
liorses taken to the Falkland Islands have, during successive 
generations, become smaller and weaker, whilst those which have 
run wild on the Pampas hate acijuirod larger and coarser heads ; 
iind such changes are manifestly due, not to any one pair, but to 
all the individuals having been aubjeetcd to the same conditions, 
aided, perhaps, by the principle of reversion. Tho new sut>4 
breeds in such cases aio not descended from any single pair, butt 
from many individuals which have varied in difierent degrees,! 
but in the same general manner; and we may conclude that thet 
races of man have been similarly produced, the modificationdi 
being either the direct result of exposure to different conditions,! 
or the indirect result of some form of selection. But to this 
latter subject we shall presently rfetum. 

On the Esctindion of Uie Saces of Mart. — The partial or eomplcta 
extinction of many races and sub-racea of man is historically 
known. Humboldt saw in South America a parrot which was 
the sole living creature that could speak a word ot the language 
of a lost tribe. Ancient monuments and atone iraplementa 
found in all partsof the world, about which no tradition has been 
preserved by the present inhabitants, indicate much extinction. 
Some small and broken tribes, remnants of fonner races, still 
survive in isolated aad generally mouniainona districts. In 
Europe the ancient races were all, according to Sehaaffhausen,'^ 
" lower in the scale than (he rude* bving savages ;" they must 
therefore have difBa-ed, to a certain estent, from any existing 
raoa. The i'emaius described by Professor Broca from Les Eyzies, 
though they unfortunately appear to have belonged to a sit^la 
family, indicate a race with a most singular combination of low 
or simious, and of high characteristics. This race is "entirely 

• Tcaaslalion in 'Aatliripoli^ical Review, Oct. 1888, p. *3l 


Tne Descent of Man. 

" different from any other, ancient or modem, that we have ever 
"heard of."" It differed, therefore, from the quaternary race of 
the cavcma of Belgium. 

I Man can long resist coaditioiis which appear extremely ua- 
favociable for his existence," He has loi^ lived in the extreme 
regions of the North, with no wood for his canoes or implements, 
and with only blubber as fuel, and meltfid snow as drink. In 
the sonthern estremiiy of America the Fuegiana survive with- 
out the protection of clothes, or of any building worthy to be 
called a hovel. In South Africa the aborigines wander over arid 
plains, where dangerous beasts abound. Man can withstand the 
deadly influence of the Terai at the foot of the Himalaya, and 
the pestilential shores of tropical Africa. 

I Extinction follows chiefly from the competition of tribe with 
I tribe, and race with race. Various checks are always in action, 
serving to keep down the nttmbers of each savage tribe, — such 
as periodical famines, nomadic habits and the consequent deatla 
of infants, prolonged sucklii^, wars, accidents, sickness, licen- 
tiousness, the stealing of women, infanticide, and especially 
lessened fertility. If any one of these checks increases in power, 
even slightly, the tribe thus affected tends to decrease ; and 
when of two adjoining tribes one becomes less numerous and less 
powerful than the other, tho contest is soon settled by war, 
slaughter, cannibalism, slavery, and absorption. Even when a 
weaker tribe is not thus abruptly swept away, if it once begins 
to decrease, it generally goes on decreasing until it becomes 

When civilised nations come into contact with barbarians the 
struggle is short, except where a deadly climate gives its aid to 
tho native race. Of tho causes which lead to the victory of 
civilised nations, some are plain and simple, others complex and 
obscure. Wo can sec that the cultivation of the land will 
be fetal in many ways to savages, for they cannot, or will not, 
change their habits. New diseases and vices have is some cases 
proved highly destructive; and it appears that a new disease 
often causes much death, until those who are most susceptible 
to its destructive influence are gradually weeded out ;" and so it 
may be with the evil effects from spirituous liquors, as well aa 
■witti the unconquerably strong tast« for them shewn b j so many 

» 'Tiansact. InterDst. Coagress 

ttrbeo der Natnrvijlker,' 1868, . 

(f Prehistoric Arch,' 1868, pp. 172- 

« Gerland(ibid.8. 12)Bive3 

175. See also Brosa (translation) 

in support of tUi statement. 

» See remarta to this effec 

!888, p. 410. 

Sir H. Holljmd's' Medical SoUl 

>i Dr. GerlaB.i ' Ueb*t das Aum- 

Reflections,' 183S, p. 3S0. 


Chap. VTI. TJu Exthiction of Races. 183 

Bavagt's. It fuither appears, mystorioua as is the fad that 
the first meeting of distinct and separated people genetatea 
diKtasa" Mr. Sproat, who in Vancotiver Island closely attended 
tu tho sulijecl; of extiactioti, beiieTed that changed habits of life, 
CL>nt>equent on the advent of Europeans, induces much ill health. 
Ue Iaj%, alGO, great stress on the apparently triSing (?auso that 
the ttatives become " bewildered and dull by the new life around 
" them ; they lose the motives for exertion, and get no -^w ones 
" in their place."" 

Tho grade of their civilisation seems to be a most important 
element in the success of competing nations. A few centuries 
ago Europe feared the inroads of Eastern barbarians ; now any 
such fear would be ridiculous. It is a more curious fact, as 
Mr. Bagehot has remarked, that savages did not fonnerly waste 
awaj before the classical nations, as they now do before modem 
civilised nations; had Ihey done so, the old moraUsts would 
have mused over the event; but there is no lament in any writer 
of that period over the perishii^ barbarians." The most po-i 
tent of all the causes of extinction, appears in many cases fo be 
lessened fertility and ill-health, especially amongst the children,) 
arising from changed conditions of life, notwithatanding that the 
new conditions may not he injuiioua in themselves. I am much 
indebted to Mr. H. H. Howorth for having called my attention to 
this subject, and for having given me information respecting it 
I have collected the following cases. 

When Tasmania was first co'lonised the natives were roughly 
estimated by somo at 7O0O and by others at 20,000. Their 
number was soon greatly reduced, chiefly by fighting with the 
English and with each other. After the famous hunt by all the 
colonists, when the remaining natives delivered themselves u|> 
to the government, they consisted only of 120 individuals," 
who were in 1832 transported to Flinders Island. This island, 
situated between Tasmania and Australia, is forty miles long, 
and from twelve to eighteen miles broad: it seems healthy, 
and the natives were well treated. Nevertheless, they suffered 
greatly in health. In 1834 they conaisted (Bonwick, p. 250) of 
forty-seven adult males, forty-eight adult females, and sixteen 
children, or in all of 111 souls. In 1835 only one hundred were left. 

» I have coll8ct«d (' Journal of Savaje life,' 1868, p. 284, 
Ee5airclna,Vojageofthe"Beagle,"' " Bngehot, ' Phyries and Poli- 

p. 4a5) s good many cases beannc; tics,' ' Fortnightly Keview,' April 
nn thi> EUbjectj lee also GcrJaod, 1, ]868, p. Vo'o. 
ibid. 9. 8. Poeppig speaks of the " All the statemants licre Rives 

"breslh of ciTiiiaalion as po 
« to savages." 

'* Sjiroat, * Scenes ttnil Stt 


iS4 ^-^^ Descent of Man. Pabt l 

As thoy continueil rapidly to deoreaso, and as tlwy themselvea 
thoughttliattheystauldiiot perish soquicMj elsewhere, they weia 
romoved in 1847 to Oyster Cove in the southempart of Tasmania. 
They then consisted (Deo. 20th, 1847) of fourtoea men, twenty- 
two women and ten children.'" But the change of Kite did no good. 
Disease and death still pursued them, and iu 186i one man (who 
died m ia69), and three elderly women alone snrrived. The 
infertility of the women is even a more remarkable fact than 
the hability of all to ill-health and death. At the time when 
only mue women were left at Oyster Cove, they told Mr. Bonwiok 
(p 3a6), that only two had ever borne children : and these two 
had ti^ether produced only three childicn \ 

With respect tothe cause of tliis extraordinary state of thinKJ!, 
Dr. Story remarks that death followed the attempts to civilLso 
the natives. " If left to fhemselves to roam as they were wont 
" and undisturbed, they would have reared more children, ond 
■' there would have been less mortality." Another carefnl 
obserrer of the natives, Mr, Davis, remarks, " The births have 
" been few and the deaths numerous. This may have been in a 
" great measure owing to their change of living and food ; but 
" more so to their banishment from the mainland of Van Diemen's 
"Land, and consequent depression of spirits" (Boawick, pp. 

a88, sao). 

Similar facts have been observed in two widely difforont 
parts of Australia. The celebrated explorer. Mr. Gregory, told 
Mr. Bonwick, that in Queensland " tho want of reproduction 
"was being already felt with the blacks, even in the most 
" recently settled parts, and that decay would set in." Of 
thirteen aborigines from Shark's Bay who visited Murohison 
River, twelve died of consumption within three months." 

The decrease of the Maories of Now Zealand has been carefully 
investigated by Mr. Penton, in an admirable Eeport, from which 
all the following statements, with ono exception, are taken." 
The decrease in number since 1830 is admitted by every one, 
including the natives themselves, and is still steadily progress- 
ing. Although it has hitherto been found impossible to take an 
actual census of the natives, their numbers were carefully 
estimated by residents in many districts. The result seems 
trustworthy, and shows that during the fourteen years, previous 

id the ■ Last of the 

■• This is tho staleraeDt of the 

1870, p. 90; an 

flovoriiur of Tasamnin, Sir W. Dani- 

Tosmapiaas,' 18^ 

,oa, ' Vsrieties of Vice-Kegal Life,' 

" 'Observalio 

1870, ToL i. p. 87. 

Inhabitants of 1 

" Fot these casej, nee Boowick'a 

iished by the Gj 

'Daily Ufe of the l'a«mnn 


Cbaf. VII. The Extimtion of Races. 185 

to 1858, the decrease was 19.42 per cent. Some of the tribes, 
thus cftrefttllj osamined, lived above a linndred miles apart, 
Home on the coast, some inland ; ojid their Tueans of subsistenoo 
and habits differed to a certain extent (p. 28). The total 
liamber in 1858 was beheved to be 53.700, and in 1872, after a 
second interral of fourteen years, another census was taken, 
and the number is given as only S6y')59, shewing a decrease ol 
32-29 per cental" Mr. Fenton, after shewing in detail the in- 
sufficiency of the various causes, usually assigEed in explana- 
tion of this estraordinary decrease, such as new diseases, the 
profl^cy of the women, drunkenness, wars, &o., concludes on 
weighty grounds that it depends chiofiy on the unproductiveness 
of the women, and on the extraordinary mortality of the young 
children (pp. 81, 84). In proof of this he shews (p. 33) that in 
1844 there was one non-adult for every 2'57 adults ; whereas in 
1858 there was only one non-adult for evej? 3'27 adults. The 
mortality of the adults is also great. He adduces as a farther 
causeof the decrease the inequality of thosexes; tor fewer female 
are born than males. To this latter point, depending perhaps 
on a widely distinct cause, I shall return in a future chapter. 
Mr. Penton contrasts with astonishment the decrease in New 
Zealand with the increase in Ireland; eonntriee not very dis- 
similar in climate, and where the inhabitants now follow nearly 
similar habits. The Maori(» themselves (p. 35) " attribute their 
" decadence, in some measure, to the introduction of new food 
"and clothing, and the attendant change of habits;" and it will 
be seen, when we consider the influence of changed conditions 
on fertility, that they are probably right. Tlie diminution began 
between the years 1830 and 1840 ; and Mr. Tenton shews (p. 40) 
that about 1830, the art of manufacturing putrid com (maize), 
by long steeping in water, was discovered and largely practised ; 
and this proves that a change of habits was b^inning amongst 
the natives, even when Now Zealand was only thinly inhabited 
by Europeans. When I visited the Bay of Islands in 183S. 
the dress and food of the inhabitants had aJieady been much 
modified: tbey raised potatoes, maize, and other agricultural 
produce, and cschanged them for English manufactured goods 
and tobacco. 

It is evident from many statements in the life of Bishop 
Patteson," that the Uclanesians of the New IletridcB and 
noighboniing archipelagoes, suffered to an extraordinary degree 
in health, and perished in large numbers, when they were 

" 'New Zealand,' Ij Alei. KuD- C. M. YoQoge, 1874; we mor« 
ntay, 1873, j.. 47. e.«pecially vol, i. p. 530. 


e Descent, of Man. 

Pabt T. 


removed to Now Zealsod, Norfolk Island, and other salubriona 
pliMi^s, ia order lo bo edncated aa missionanes. 

The decrease of the native population of the Sandwich IsJaniis 
is as notorious as that of New Zealand. It has boon roughly 
eBtimated by those best capable of judeing, that when Cook 
diacoYcred the Islands in 1779, the population amounted to 
about 300,000. According to a loose census in 1823, the 
nimibers thea were 142,050. In 1832, and at several subsequent 
periods, on accurate census was officially taken, but I have 
been able to obtain only the following returns : 


We hereseethatin the interval of forty years, between 1832 and 
1872, the population has decreased no less than siity-eight per 
cent.! This hasbeenattributedbymostwriters to the profligacy 
of the women, to former bloody wars, and to the severe labour 
imposed on conquered tribes and to newly introduced diseases, 
which have been on several occasions eitremeJy destructjve. No 
doubt these and other such causes have been highly efficient, 
and may account for the extraordinary rate of decrease between 
the years 1832 and 1836 ; but the most potent of all the causes 
seoTHS to te lessened fertility. According to Dr. Euschenberger 
of the U.S. Navy, who visited th^e islands between 1835 and 
1837, in one district of Hawaii, only twenty-five men outoril34, 
and in another district only ton out of 637, had a family with as 
many as three children. Of eighty married women, only thirty- 
nine had ever borne children; and " the official report gives an 
"average of half a child to each married ooiiple in the whole 


Chap. VII. TIu Extinction of Races. 187 

" ieland." This is almost esactly the same average as with tha 
Tasmanians at Oyster Ctove. Jarves, who published his History 
ia 1843, says that " families who have thiee cMldien are freed from 
" all teles ; those having more, are rewarded hy gifts of land and 
"other ettconragementB." This unparalleled enactment by the 
government well shews how infcrtOe the tace had become. The 
Itev. A. Bishop statedin the Hawaiian 'Spectefor'in 1839, that a 
iatge proportion of the children die at early ages, and Bishop 
Staley informs me that this is still the case, jiKt as in New 
Zealand. This has been aUribnted to the neglect of the children 
by the women, bnt it is probably in large part due to innate weak- 
ness of coQgtitntio<i in the children, in relation to the lessened 

fertility of their parents. There is, moreover, a further resan- 

blanc© to the case of New Zealand, in the feet that there is a 
lai^ excess of male over female births : the census of 1872 
gives 81,650 males to 25,247 females of aU ages, that is 125-86 
males for every 100 females; whereas in all civihsed coontriea 
the females exceed the males. No doubt the profligaey of the 
women may in part acconnt for their small fertility ; but their 
changed habits of life is a much more probable canso, and which 
will at the same time account for" the increased mortaUty, 
especially of the children. The islands were visited by Cook in 
1779, by Vancouver in 1794, and often subsequently by whalers. 
In 1819 missionaries arrived, and found that idolatry had been 
already abohshed, and other changes effected by the king. After 
this period there was a rapid change in almost all the habits of 
life of the natives, and they soon became " the most civilised of 
" the Pacific Islanders." One of my informants, Mr. Coon, who 
was born on the islands, remaiks that the natives have undeigono 
a greater change in their habits of life in the eonrae of fifty years 
than Englishman during a thousand years. From information 
received from Bishop Staley, it does not appear that the 
poorer classes have ever much changed their diet, although 
many new kinds of fruit have been introduced, and the sugar- 
cane is in universal use. Owing, however, to their passion for 
imitating Europeans, they altered their manner of dressing at 
an early period, and the use of alcohoUc drinks became very 
general. Althoi^h these changes appear inconsiderable, I can 
well believe, from what is known with respect to animals, that 
they might suffice to lessen the fertility of the natives," 

" The foiTgoing statements are Islands,' 1851, p. 277. Kuschen- 

taken ehiefly from the folIowiDg berger is cjooted by Bonwiek, * Lart 

works: 'JaiTes' HisWry of the of the TasmaniaBs.' 1870, p. 378. 

Hawaiian liknda,' 1843, p. 400-407. Bishop is qooted by Sir E. Belcher, 

Cheerar, ' IJfe inths Sandwich 'Voyage Konnd the World,' 184», 


tSS The Descent of Man. p*bt L 

Lastly, Sir. llacuamara states" tiat Ihe low and degraded 
inliabitants of tho Acdaioaa Islands, on ths eastern Bide of tlw, 
Gulf of Bei^al, axe " eminently susceptible lo any change of 
" climate ; in fact, take them away from their island homes, and 
" thoy are almost certain to die, and that independently of diet 
" or extrauaous inliueuces." He fntther states that tho inbabit- 
onta of tho Valley of Nopftl, which is extremely hot in suiiimor, 
and also the Tarious hill-tribes of India, suffer from dyioiiterj 
and fever when on the plaiiis; and they die if thoy attempt to 
pass tho whole year tliere. 

We thus BOO that many of the wilder races of man are apt to 

suffer much in health when subjected to changed conditions 

or habits of life, and not esclusively from being transported to 

a new climate. Mere alterations in habits, whicli do not appear 

injurious in themselTes, seem to have this game eEfect ; and in 

several cases the children are partioulai'ly liable to suffer. It 

has often been said, as Mr. Macnamara remarks, that man can 

■ resist with impunity the greatest diversities of climate and other 

I changes; but this is true only of the civilised races. Man iu 

I his wild condition seems to be in this respect nbnost ns sua- 

I cepttblo as his nearest allies, tho anthropoid apes, which have 

I never yet survived loi^, when removed from their native 

V country. 

Lessened fertility from changed conditions, as in the case of the 
Tasmanians, Maories, Sandwich Islanders, and apparently the 
Australians, is still more interesting than tlioir liability to 
ill-health and death ; for even a slight degree of infertility, 
combined with those other canses which lend to check the 
increase of every population, would sooner or later lead to 
extinction. The diminution of fertility may be explained in 
some cases by the profligacy of the women (as until lately with 
the Tahitians), but Mr. Fenton bus shewn that this explanation 
by no means suffices with tJie New Zealanders, nor docs it with 
the Tasmanians. 

In the paper above quoted, Mr. Macnamara gives reasons for 
believing that the inhabitants of districts subject to malaria are 
apt to be sterile; but this cannot apply in several of the abovii 
cases. Some writers have snggosted that the aborigines of 
islanda have suffered in fertility and health from long continued 

I have 


Chap. VIT. The Extinction of Races. 

inter-breodii^ ; but in the abow cases infertility liaa coincided 
too closely with the arrival of Europeans for us to admit this 
explftufttion. Nor have we at present any reason to believe 
that man is h%lily sensitive to the evil effects of inter-breeding, 
especially in areas so large as New Zealand, and the Sandwich 
archipelago with its diversified stations. On the contrary, it is 
knoivn that the present inhabitants of Norfolk Island are nearly 
all cousins or near relations, as are the Todas in India, and the 
inhabitants of some of the Western Islands of Sootlaud; and 
yet they seem not to have snffered in fertility." 

A much Kore probable view is suggested by the analc^ of 
the lower animals. The reproductive system, can be shewn lo be 
Buscentible to an extraordinary degree (though whj we know 
not) to changed conditions of life-, oEd this susceptibiUty leads 
both to beneficial and to evil results. A large collection of facts 
on this subject is given in chap, sviii. of vol. ii. of my 'Variation 
of Animals and Plants under Bomestication,' I can here give only 
tho briefest abstract; and every one interested in the subject 
may consult the above work. Very slight changes increase the 
healti, vigour aid fertility of most or all organic beings, 
whilst other changes aie known to render a large number of 
animals sterile. One of the most familiar cases, is that of tamed 
elephants not breeding in India; though they often breed in 
Ava, where the females are allowed to roam about the forests to 
some extent, and are thus placed under more natural conditions. 
The case of various American monkeys, both soses of which 
have been kept for many years leather in their own countries, 
and yet have very rarely or never bred, is a more apposite in- 
stance, because of their relationship to man. It is remarkable 
how sUght a change in the conditions often induces sterility in a 
wild animal when captured ; and this is the more strange as all 
our domesticated animals have become more fertile than they 
were in a stato of nature; and some of them can resist the 
mostunnaturaleonditionswith undiminished fertility." Certain 
groups of animals are much more liable than others to be 
affected by captivity ; and generally all the species of the same 
group are afi'ected in the same manner. Eut sometimes a sii^Ie 
spoeios in n group is rendered sterile, whilst the others are not 
m ; on the other hand, a single species may retain its fertility 

'* Ou the close relationship of the 
Norfolk JalaudPTS, see Sir W. Deni- 

ma, 'VHiietien of Vit»-Regal Life,' joo^.. 

vol. i. 13T0, p. 410. For the Todaa, •■ For the evidence on thb head, 

ne Col. Maishall't work, 1873, p. see 'Variation of Ai^bIb' &u.. 

110. For the Wettem Islands t{ vol. iL y. HI. 


IQO Tite Descent cf Man. Past L 

whilst most of the others fail to breed. Tlio males and femaloa 
of some species when confined, or when allowed to live almost 
but not quite free, in theic natiTo countrr, nQTcr unite; others 
thus circumstanced frequently unite but never produce oflspricg ; 
others again produce some offepring, but fewer than in a state 
of nature ; and as bearing on the above cases of man, it is 
important to remark that the young are apt to be weak and 
sickly, or malformed, and to perish at an early age. 

Seeing how general is this law of the suBceptibiUty of the 
reproductive system to changed conditions of life, and that it 
holds good with our nearest allies, the Qnadmmaiia, I can 
hardly doubt that it applies to man in his primeval state. Hence 
if savages of any race are induced suddenly to change their 
habits of life, they become more or less sterile, and their young 
offsprii^ the same manner and from the same 
cauBe, as do the elephant and htmting-leox>atd in India, many 
monkeys in America, and a host of animals of all kinds, on removal 
from their natiirai conditions. 

We can see why it is that aborigines, who have long inha- 
bitud islands, and who must have been long exposed to nearly 
uniform conditions, should bo specially affected by any change 
in their habits, as seems to be the case. Civihsed races can 
certainly resist changes of all kinds far better than savages ; 
and in this respect they resemble domesticated animals, for 
though the ktter sometimes suffer in health (for instance 
European dogs in India), yet they are rarely rendered sterile, 
liiough a few such instances have been recorded." The 
J immunity of civilised races and domesticated animals is 
probably duo to their having been subjected to a greater extent, 
and therefore having grown somewhat more accustomed, to 
diversified or varying conditions, than the majority of wild 
animals; and to their having formerly immigrated or been 
tarried from country to country, and to different families or 
sub-races having inter-crossed. It appears that a cross ivith 
civilised races at once gives to an aboriginal race an immunity 
from the evil consequences of changed conditions. Thus the 
crossed offspring from the Tahitians and English, when settled 
in Pitcairn Island, increased so rapidly that the island was soon 
overstocked; and in June 1856 they wore removed te Norfolk 
Isiand. They then consisted of 60 married persons and 134 
Bhildren, making a total of 194. Here they likewise in- 
creased so rapidly, that although sixteen of them returned to Pit- 
caim Island in 1859, they numbered in January 1868, SOOsoulSj 

" 'Variation of Animal*,' Ac, vol. iU p 16. 


Ohap. VIL The Extinction of Races. 191 

the maJes and femalea being in. exactly equal numUire. What a 
contrast does tliis case present witt that of tho Tasmanians ; 
the Norfolk Islanders ina-eited in only twelve and a half years 
from 194 fo aOO; whereas the Tasmanians (k<.reti'<(d during 
fifteen years from 120 to 46, of which latter number only ten 
were children." 

So again in tho interval between tie census of ISbO and 1872 
the natives of full blood in the Sandwich Islands decreased by 
8081, whilst the half-castes, who are believed to be healthier, m- 
ereased by 847 ; but I do not know whether tho latter number 
includes th^ofispring from tho half-castes, or only the half-castes 
of the first generation. 

The cases which I have here given all relate lo aborigines, 
who have been Bubjeoted lo new conditions as the result of tho 
immigration of civilised men. But sterility and ill-health would 
probably follow, if savages were compellai by any cause, such 
OB the inroad of a conquering tribe, to desert their homus and 
to change their habits. It is an int^restii^ ciccnmstance that 
the chief check to wild animals becoming domesticated, which 
implies the power of their breeding freely when first captured, 
and one chief check to wild men, when brought into contact 
with civilisation, surviving to form a civilised race, is the same, 
namely, sterility from changed conditions of life. 

Finally, although the gradual decrease and ultimate extinction 
of the races of man is a highly complex problem, depending on 
many canses which differ in different places and at difibrent 
times; it is the same problem as that presented by the extinc- 
tion of one of the higher animals — of the fossi! horse, for in- 
BtaccOj which disappeared from South America, soon afterwards 
to bo replaced, within the same districts, by counties troops 1 

of the Spanish horse. The Kew Zealauder seems conscious of --^ 
this parallelism, for he compares his futiHC fate with that of ^ 
the native rat now ahnost exterminated by the European rat, j" V* / ' 
Though the diffieulty is great to our imagination, and really ^/i •; ' 
great, if we wish to ascertain the precise causes and their \y ,' 
manner of action, it ot^ht not to be so to our reason, as long as •f' Z' 'y 
we keep steadily in mind that tho inci'easo of each species and 
each race is constantly chocked in various ways ; so that if any 
new check, even a sl%ht one, be superadded, the race will surely \ 
decrease in number; and decreasing numbers will sooner- or 

" Theao dftails are taken ftom May 39th, 1863- Tho following 

'The MHtineera of the " Bounty," ' ttatemeiits about, the Sandwich 1*- 

by Lady Belcher, 1870; and from landeia are firom the 'UoocUIb 

' PitGurn Island,' ordered to he Gaziette,' and liom Mr, Coan. 
pilni«d>y the Hoasi " " 



The Descent of Ma}u 

later load to extinction ; the end, ia mOKt onses, being promptly 
determined by the inroads of conquoring tribes. 

, On the B'oi'malioit of the Races of Man.— In some cases tho 
I crcssing of distinct races has led to the formation of a new rni^e. 
The singular fact that Europeans Eiud Hindoos, who belong lo 
the same Aryan stock, and speak a. lengoage fondamentally tlio 
same, differ widely in appearancej whilst Eurojwans differ hut 
little from Jews, who belong to the Semitic stock, and speak 
quite another language, has been aeoounted for by Broca," 
through certain Aryan branches having been lai^ely crossed 
by indigenous tribes duriug their wide diffusion. When two 
k races in close contact cross, the first resnlt is a hefcn^eneous 
' mixture: UiusMi'.Uunter, iii describing the Santali or bill-tribes 
of India, says that hundreds of imperceptible gradations may be 
traced " from the black, squat tribes of the mountains to the fall 
" olire-colonred Brahman, with his intellectual brow, calm '■yes, 
"and high but narrow head;" so that it is necessary in courts 
of instice to ask the witnesses whether tliey are Santalis or 
Hindoos." Whether a hetert^neous people, such as the inhabi- 
tants of some of the PolyceEian islands, formed by the crossing 
of two distinct races, with few or no pure members left, would 
ever become homc^eneous, is not known &om direct evidence. 
But as with onr domesticated animals, a cross-breed can certainly 
be fixed and made uniform by careful selection" in the course of 
a few generations, we may infer that the free intercrossing of a 
betcrc^neous misture during a long descent would supply the 
place of selection, and overcome any tendency to reversion ; so 
that the crossed raca would ultimately become homogeneous, 
though it might not partake in an equal degree of the cliaraeters 
of the two pai'ent-races. 

Of all the differences between the races of man, the colour ol 
the skin is the most conspicuous and one of the best marked. I', 
was formerly thought that differences of this kind conld be 
accounted for by long exposure to different climates; but 
I'allaa first shewed that this is not tenable, and he has since been 
tciUowed by almost all anthropologists."' This view has been 
rejected chiefly because the distribution of tha variously 
coloured races, most of whom must have long intiabited their 

'• 'Oo Anthropology,' transla- " PalJaa, ' Act. Atvi. St. Peters- 

tica ' Authropolog. Review,' Jan. burg,' 1780, part il. p. S9. He ».i3 

18BS, p. 38, followed bj Eiiilolphi, ia his ' Bcy- 

*• 'The AddsIs of Karat Bengal,' traga lar Anthiopologie,' mVi, 

.868, p. 134. An eicellent anraraar)' of the oyi- 

" 'Tha Variation of Animals ajjd deuce is givan by Godion, ' Dt 

PlanM under Domeslicr.tion,' vol. ii. PEspiee,' 1859, vol. ii. p. 348, Sit 


Chai'. VII. The PcriHatiott of Races. 193 

present homes, does not coincide with corresponding diftdrenceS 
of climate. Some little weight may be given to sueh caseB as 
that of the Batch families, who, as we hear on excellent autto- 
rity," have not undergone the least change of colonr after 
residing for three centuries m South Africa. An ailment on 
the same side maj hkewise be drawn from the uniform appear- 
ance in Tarioua parts of the world of gipsies and Jews, though 
the uniformity of the latter lias been soraewhat exaggerated." 
A Teiy damp or a very dry atmosphere has been supposed to bo 
more influential in moditying the colour of the skin than mere 
boat ; but as D'Orbigny in Soutli America, and Livingstone in 
Africa, arrived at diametrically opposite conclusions with respect 
to dampness and dryness, any conclusion on this head must bo 
considered as very doubtful." 

Various fa«ts, which I have given elsewhere, prove that the 
eoloni of the skin and hair is sometimes correlated in a surpris- 
ing manner with a complete immunity from the action of certain 
vegetable poisons, and from the attacks of certain parasites. 
Hence it occurred to me, that negroes and other dark Kuses 
might have acquired their dark tints by the darker individuals 
escaping from the deadly influence of the miasma of their 
native countries, during a long series of generations. 

I afterwards found that this same idea had long ago occurred 
to Dr. Wells. "• It has long been known that negroes, and even, 
mulattoes, are almost completely exempt from the yellow -fever, 1 
so destructive in tropical America.*' They likewise escape to a t 
lai^ extent the fatal intermittent fevers, that prevail along at 
least 2600 miles of the shores of Africa, and whiuh annually 
cause one-fifth of the white settlers to die, and another fifth to 
return home invalided.*' This immunity in the negro seems to 
be partly inherent, depending on some unknown peculiarity of 
constitution, and partly the result of acclimatisation. Pouchet" 

" Sir Andrew Smith, as quoted the Historical Sketch <p. ivi.) to my 

It Knoi, ' Rates of Man ' 1850, p. ' Origin of Species.' Various caa^s 

4-73. of colcur co.relatod with constitu- 

" Sec De Quatrefoges on this tionsi peouliacitiei are given in my 

head, 'Rbvuo des Cours Scieati- 'Vaiiation of Animals under Do- 

fiqoes,' OcL 17, 1863, n. 731. meBticatloo,' vol. ii. pp. 237, 335. 

" Livingstone's 'Travels Hod Re- "See, for inst-Qoe, Nott and 

Marches in S. Africa,' 1857, pp. Gliddon, 'Types of Mankind,' p. 68. 

338, 329. D'Orbigny, as rjuoted liy " Major Tulloeh, In a paper I'eart 

Godron, 'Do I'Espico,' vol. ii, p. before the StatiBtical Society, Aprt: 

2tie. 21}th, IS40, and gliea in the 'Athe. 

■• See a paper read before the nicum,' 1840, p. 353. 

Eoval Soc. ID 1813, and pnblished " 'The Flurolity of the Huni.ii 

In Cih.Essays in 1318. I have given Race' (ti^anslat.), 1864, p. 60. 
ui account of Dr. Wells' vie^vs in 


194 The Descent of Man. ^.-..w-i I. 

Etatea that the negro resimenta recrnitod near the Soudan, and 
Borrowed from the Viceroy of Egypt for the Mexican war, 
escaped the yellow-fever almost equally with the negroes origin- 
ally brought ftomtarioua parts of Africa aod accustomed to the 
tlimate of the West ladies. That a«climatisation plays a part, 
is shewn hy the many cases in which negroes have become some- 
what liaUe to tropical fevers, after having resided for some time 
in a colder climate,"" Thenatureof the climate under which the 
white races have long resided, likewise has some inflneuce on 
them ; for during the fearful epidemic of yellow-fever in 
Demerara during 1837, Dr. Blair found that the death-rate of the 
immigvants was proportional to the latitude of the country 
whence they had come. With the negro tie immunity, as fer as 
it is the result of acclimatisation, implies exposure during a 
prodigious length of time ; for the aborigines oE tropical America 
who have resided there from time immemorial, are not exempt 
from yellow fever; and the Eev. H. B. Tristram states, that 
there are districts in Northern Africa which the native inhabit- 
ants are compelled annually to leave, though the negroes can 
remain with safety. 

That the immunity of the negro is in any degree correlated 
■with the colour of his skin is a mere conjecture : it may be 
[Correlated with some difference in his blood, nervous system, or 
|other tissues. Nevertheless, from thefticfs above alluded to, and 
from some connection apparently existing between complexion 
and a tendency to consumption, the conjecture seemed to me 
not improbable. Consequently I endeavoured, with but little 
success," to ascertain how iar it holds good. The late Dr. 

" Quatrefages,' Unite del'Eapice " Is some limited deBres of rd.ition 
H ■ ' 18-1 "05 W ■ ' b races of 

" onr donicstio animalaof a relation " aad light-coloured liair, and Laii 

" between the colour of the dermal " of iat«miediato or doubtful tints ; 

" appendages and the constitntion ; "wit jf j similar account were 

■ iB& it being dotorioas that thwe " fcent by It* same medicaj (;™tl*- 


CFir, VII. 

The Formation of Races. 


Daniel], who had long lived on the West Coast of Africa, told me 
that he did not belieye in any such relation. He was himself 
miusually foir, and had withstood the climate in a wonderfnl 
manner. When he first anived as a boj on the coast, an old and 
experienced negro chief predicted from his appearance that this 
would prove the case. Dr. Micholeon, of Antigna, after haTing 
attended to this euhject, writes to ma that he does not think that 
dark-coloured Bnropeans escape tlie yellow-fever more than 
those that are Ught-colonred. Mr. J. M. Harris altogether 
denies that Entopeans with dark hair withstand a hot climate 
better than other men : on the contrary, experience has taught 
him in making a selection of men for service on the coast of 
Africa, to choose those with red hair."' As far, therefore, as 
these slight indications go, there seems no foundation for the 1 
hypothesis, that blackness has resulted from the darker and \ 
darker individuals having survived better during loi^ exposure ) 

Dr. Sharps remarks," that a tropical sun, which bums and 
blisters a white skin, does not icjnre a black one at all ; and, as 
he adds, this is not due to habit in the individual, for children 
only six or eight months old are often carried about naked, and 
are not affected. I have been assured by a medical man, that 
some years ago during each summer, but not during the winter, 
his hands became marked with ligM brown patelics, like. 

" of generations." 

" ' Anthropoli^ical Review,' Jan, 
18G6, p. iii. Dr. Sharpe also says, 
with respect to India (' Man a Spe- 
laal Creation," 1873, p. 118), that 
" it has been noticed fay some medi- 
" caJ officers tliat Europeans with 
" light hair aud florid complexions 
" BUlfer leas IVom di .... 

, of all the men who enflered 

n dys. . . 
" be apparent, after some thonsand 
" cases had been tabulated, whether 
" there exists aor relation between 
" the colour of the hair and cansti' 
" ijutional liability to tropical dis- 
" eases. Perhaps no such relation 
" would be discovered, but the in- 
" vestigation is well worth making. 

■' hair 

with dark 

■esult would be of high 

i iudica 


" by which 
" from a remote period an un- 
" healthy tropical climate, might 
" have becoraa dark-coloured fay 
" ths better preservation of dark- 
" haired or dart-compleiloned in- 
' diriduals daring a long succession 

" remark." On the other band, 
Mr. Heddle, of Sieri'a Leone " who 
" has had more clerks killed under 
" him than any other man," by tha 
climate of the West African Coast 
(W. Beade, 'African Sketch Book,' 
vol. ii. p. 532), holda a directly 
opposite view, as does Capt. Barton. 
" 'ManaSpeoia]Creation,'ia73 
p. 119. 



196 The Descent of Misn. Pa bt L 

fttthongh lai^Br than freckles, and timt these patches were nevta 
affected by sim-buxning, whilst tbe white parts of his skin 
have on KCTeral occasions been much inflamed and blistered. 
With the lower animals there is, also, a constitutional difference 
in liabilitj to the action of the Bun between those parts of the 
skin cloljied with white hair and other parts." "Whether the 

I saving of the skin from being thus burnt is of sufficient impor- 
tance to aceoimt for a dart tint having been gradually acqtnred 

I by man through natural selection, I am unable to judge. If it 
be so, we should have to assume that the natives of tropical 
America have lived there for a much shorter time than tho 
negroes in Africa, or the Papuans in the southern parts of the 
Blalay archipelago, just as the lighter-colonred ffindoos have 
resided in India for a shorter time than the darker aborigines of 
the central and Bonthem parts of the peninsula. 

Although witlronr present knowle<^^6 cannot accoimt for 
the differences of colour in the races of man, through any 
advantage thus gained, or from the direct action of climate ; yet 
we must not qnite ignore the latter agency, for there is good 
reason to believe that some inherited effect is thus produced^ 

We have seen in the second chapter that the conditions of life 
affect the development of the bodily frame in a direct manner, 
and that the effects are transmitted. Thus, as is generally 
admitted, the European settlers in the United States undergo u 
slight but extraordinarily rapid change of appearance. Thei? 
bodies and limbs become elongated ; and I hear from Co/. 
Bemys that during the late war in tbe United States, good 
evidence was afforded of this fact by the ridiculous appearance 
presented by the German regiments, when dressed in ready-maao 
clothes manufactured for the American market, and which were 
much bio long for the men in every way. There is, also, a con- 
siderable body of evidence shewing that in the Southern States 
tbe honse-slaves of the third generation present a markedly 
different appearance from the field-slaves.'' 

" Variation of Animn'j nnd settlod in GeorgEa, havs »oqnlr*d in 

riant! under Domestication,' toI. it tha coui^ae of two generationn dalle 

pp. 336, 337. hair and eyas. Mr. D. Forbes ia- 

" See, for instance, Quatreftgcs forms me that the Quidioaa in iBo 

(' EevuB des Coura Scleatifiqnes,' Andes varv greatly iu cxilour, hl- 

Oct. 10, 1868, p. 72*) on the effeeia cnrding m 'the position of the ralleya 

3f residence inAbyaaiuia and Arabia, inhabited by tliem. 
-i.nd otlier analogous cases. Ih>. " Harlan, 'Jledical Keseatches.' 

itvlle (' Der Menscb, sejus Abstain- p. 533. Quatietages (' Units' de 

mong,' &c., ]8e5, s. 99) states, on I'Esptce Huinaine,' 1861, p. 128) 

the authority of Khanikof, that the has collected much evidence on Ihii 


CffAP, vn. The Formation of Races. 197 

If, however, we look to the races of man as distrihuted over 
the world, we must tafer that their characteristic differenees can- 
not be accounted for by the direct action of different conditions 
of life, even after exposure to them for an enormous period of 
time. The Esquimaux lire eselusively on animal food ; they are 
clotted in thick fur, and a^e exposed to intense cold and to 
prolonged darkness; yet they do not differ in any extreme 
degree from the inhabitants of Southern China, wto live entirely 
on vegetable food, and are exposed almost naked to a hot, glaring 
climate. The unclothed Foogians live on the marine produc- 
tions of their inhospitable shoi'es; the Botoeados of Brazil 
wander about the hot forests of the interior and live chiefly on 
vegetable productions ; yet these tribes resemble each other so 
clcsely that the Fuegiaus on board the " Beagle " were mistaken 
by some Brazilians for Botocndos. The Botooudos again, as 
well OS the other inhahitanta of tropical America, arc wholly 
different from the Hegroes who inhabit the opposite shores of 
the Atlantic, are exposed to a nearly similar climate, and follow 
nearly the same habits of life. 

Nor can the differences between the races of man be accoimted \ -) 
for by the inherited effects of the increased or decreased use of 1 
parts, except to a quite insigniflcant degree. Men who habitu- j 
ally live in canoes, may have their l^s somewhat stunted ;/ 
those who inhabit lofty regions may have their chests enlai^d ; 
and those who constantly cse certain sense-organs may have the 
cavities in which they are lodged somewhat increased in size, and 
their featnres consequently a little modified. With civilised 
nations, the reduced size of the jaws from lessened use — the 
habitual play of different muscles serving to express different 
emotions — and the increased size of the brain from greater 
intellectual activity, have together produced a considerable 
effect on their general appearance when compared with 
Kavages._^ Increased bodily stature, without any corresponding 
increase in the size of the brain, may (judging from the pre- 
viously adduced case of rabbits), have giren to some races an 
elongated skull of the doUchooeplialic type. 

Lastly, the little-understood principle of con'elated develop- 
ment has sometimes come into action, as in the case of great 
muscular development and strongly projecting supra-orbital ' 
ridges. The colour of the skin and hair are plainly correlated, as ' 
is the texture of the hair with its colour in the Mandans of 
North America.'* The colour also of the skin, and tho odour 

" Mr. CatUn states ('N, Ameri- 
ean Indiana,' 3rd edit. 1842, vol. I. 
p. 49) that in the whole trib* of 


1 98 The Descent of Man. Past i. 

emitted Tiy it, are likewise in some manner connocted. With tiie 
breeds of sheep the number of hairs within a given space and the 
nwmher of the excretory pores are related." If wo may judge 
from the analogy of our domesticated animals, many modifica- 
tions of structure in man probably come under this principle of 
correlated development. 

We baye now seen that the external characteristic differences 
between the races of man cannot be accounted for in a Eatisfac- 
tory manner by the direct action of the conditions of life, nor by 
the effects of the continued use of parts, nor through the 
principle of correlation. '\Vb nr^i therefore led t o inqu ire 
whether slig ht individual differences , Xq "tiii ;jh tob"' is emi- 
nently habi e, m ay not have been preserved and augmented 
during a long senea ot 'g^erations tniongh-Satura l .seJoctioB, . 
Jiut here we are at once met by the objection that beneficial 
variations alone can be thus preserved ; and as far as we are 
enabled to judge, although always liable to err on this head, none 
of the differences between the races of man are of any direct or 
special service to him. The intellectual and moral or social 
faculties must of course be excepted from this remark. The great 
■variability of all the external differences between the races of man, 
likewise indicates that they cannot be of much importance ; for 
if important, they would long ago have been either fixed and 
preserved, or eliminated. In this respect man resembles those 
forms, called by naturalists protean or polymorphic, which have 

I remained extremely variable, owing, as it seems, to such varia- 
tions being of an indifferent nature, and to their having thus 
fBoaped the action of natural selection. 

We have thus far been baffled in all our attempts to account 
for the differences between the races of man ; but there remains 
one important agency, namely SesuEil Selection, which appears to 
have acted powerfully on man, as on many other animals. I do 
not intend to assert that sexual selection will account for all the 
differences between tbe races. An unexplained residuum is left, 
about which we can only say, in our ignorance, that as indivi- 
duals are continually born with, for instance, heads a little 
rounder or narrower, and with noses a little longer or shorter, 
such slight differences might become fixed and uniform, if the 

the Mandans, abont oat in tea or fioe and soft. 

tweke of the memliei^, of alt ages •' On the odonc of the ekln. 

and both seies, have bright silrery Godron, ' Sur I'Esptce,' torn. ii. |'. 

gvej hair, tvhich is hereditary. 217. On the pores in the skic, 

Now this hair is as coarse and Dr. Wilokeus, 'Die Aofgaben der 

barsh as that of a horse's mane, Landwirth. Zoolechnik,' iStfV, a. 7. 


Chap, VIL Structure of the Brain. 199 

unknown agencies whicli induced ttem were to act in a more 
constant manner, aided by long-continued intercrossing. Such 
variations come under tiia provisional class, alluded to in our 
second chapter, wMct for the want of ft better term are often 
called spontaneous. Nor do I pretend that the effects of sesual 
BclectJon can be indicated with scientific precision ; but it can be 
shewn that it would be an inexplicable fact it man hftd not been 
uiodiQed by this agency, which appears to have acted powerfully 
on innumerable animals. It can fnrther be shewn that the 1 
differences between the races of man, as in colour, hairinesB,! 
form of features, &o., are of a kind which might have beeul 
expected to come under the infiucnce of sexuaJ selection. But in 
order to treat this subject properly, I have foand it necessary to 
pass the whole animal kingdom in review. I have therefore 
devoted to it the Second Part of this work. At the close I shall 
return to man, and, after attempting to shew how far he has 
been modified thiough sesual selection, will give a brief summary 
of the chapters in this First Part, 


AND IBB Development of the Bbaih ik Man and Ares. Bi 

Phofesbob Huslev, F.K.8. 

The controveray respeoliiig the ruiture and the extent of Ihe iliffer- 
encee in the alrauture of the brain in man und the apes, whifh arose 
some flfleea years ago, has not yet come to an ecui, though thu Btibjttt 
matter of the dispute is, at present, totally differeut from what it was 
formerly. It wpa origin.iily asserted and re-asserted, with singular 
perhnacity, that the brain of all the apes, even the highest, diflcrs from ■ 
[hat of man, in the absence of BUch eoiispicuons otrnutureB aa the 
posterior lobes of the cerebral hemispheres, with the posterior comu of 
the lateral ventriele and the Jt?jipocantinw winor, coutuined in those 
lobes, which are eo obvions in man. 

But the truth that the three structures in question are a* well deve- 
loped in apes' as in human brains, or even better ; and that it la charucter- 
istie of all the Primo/es (if we exclude the Lemuig) to have these parts 
wdl developed, atauds at present on as secure a baeia as any proposition 
in comparative anatomy. Moreover, it is admitted by every one of the 
long seri«e of anatomists nho, of late years, have paid special attention to 
the arrangement of the compUcated sulci and gyri which nppear upon 
the sni'bce of the oeiebrul hemispheres in man and the higher apes, 
that they are disposed eflei the very same pattern in him, as in them. 
Every piincipnl gyros and sulcus of a chimpanzee's brain ia clearly 
represented in that of a man, ho that tlie tetminology which eppties to 
the one answers for ihe other. On this point there ia no difference of 

Zioion. Some yeers einoe. Professor Bisthoff published a memoir '" on 
e cerebittl convolutions of man and apes; and as the purpose of 
my learned colleHgoe was certainly not to diminish the value of the 


200 Tke Descent of Man. Paht L 

d ifferenoes between apes and men in this respoet, I am glad to make a 

"That theapes, nnd eapeeially the orang, ehimpanzee and gorilla, 
" come veiycioaB to man in their orgamgatiOD, muuh nearer than to any 
" other animal, ia a well known fact, diaputed by nobody. Looking at 
" the matter from the point of view of or^anisiition alone, no one probably 
" nonld ever hsTB diapntad (he view of Linmens, that roan ehould be 
" plaood, merely as a peonliar apeeiea, at the bead of the mammalia and ot 
■* those apes. Both shew, in al! their orgaiia, so close an affinity, that the 
" most exact anatomical investigation is needed in order to demonstrate 
'■ those differences whiob really exist. So it is wili lie brains. The 
" brairs of man, tbe orang, the chimpanzee, Uie gorilla, in spite of all 
■' Ike important diSerences which they present, come very close to one 
■■another" (L c. p. 101). 

Tbere remains, then, nodiapute as to the resemblance infondamental 
characters, between the apesbtainand man's; nor any as to the won- 
derfully close similarity between the ohimpanaee, orang and man, in 
even the details of the arrangement of the ^ri and suici of the cerebral 
hemispheres. Nor, taming to the differences between the brains M 
the highest apes and that of man, is there any serious question as Ic 
the nalure and eitent of these differences. It is admitted that the man's 
ctrebriil hemispheres are absolutely and relatively larger than those of 
the orang and chimpanzee : that his frontal lobes are leas escavated by 
the npwanl protrusion of the roof of the orbits ; that hia gyri and sulci 
are, as a role, lea symmetrically disposed, and present a peater irambcr 
of secondary plications. And it is admitted that, as a rule, in man, lie 
teraporo-ocoipitol or"eitenial perpendioalat''fiaanre, which is usually 
so Btronglv marked a feature of the ape's brain is but faintly marked. 
But it is also clear, that none of these difFerenees constitutes a shaip 
demarcation between the man's and the ape's brain. In respect to the 
oitemal perpendicular flssure of Gratiolet, in the hnman brain, for 
instance. Professor Turner remarks ;" 

" In some brains it appears simply as an indentation of the margin ol 
" the hemisphere, but, in others, it extends for some distance more or less 
" transversely outwards. I saw it m the right hemisphere of a female 
" brain pass moie than two inches outwards ; and in another specimen. 
" also the right hemisphere, it proceeded for fonr-tenlhe of an inch out- 
" wards, and then extended downwards, aa far as the lower margin of the 
" outer smfaee of the hemisphere. Theimperfectdeflnitionof this liasnro 
" in (he majority of human brains, as compared with its remarkable dis- 
" tioctness in the brain of most Qnadmmiuia, is owing to the presence, in 
" Ibe former, of certain superficial, well marked, secondaty eonvolutiona 
" which bridge It over and connect the parietal with the occipital lobe. 
" The closer the first of these bridging gyrl lies to the longitndinal 
" Assure, the shorter is the external parieto-ocoipilal fissure." (Lap. 12.; 

The obliteration of the external perpendicular fissure of Gratiolet, 
therefore, is not a constant character of the human brain. On the other 
band, its full development is not a constant character of the highBi 
ape's brain. For, in the chimpanzee, the more or less extensive oblitera- 
tion of the external perpendicuhtr sulcas by " bridging convolutions," ou 
one side or the olficr, has been noted over and over again by Prot 

" ' CocvolMtions Iff the Hurpwn Ceiehinni TopograrinKally Consi.lfiaa. 

'ieo,p, 18, 


Chap, VII, Structure of t/ie Brain. 20 1 

BuCestnn, Mr. MarBhall, M. Brora aod Professor Turner. At ths 
candiision of a apeoial papor on thia sntjeot the latter writes :" 

■' The tJiree speoimeiia of the brain of a chimpanzee juat deficrifced, 
" prove, that the generalisation which Gnttiolethasatlempted to draw of 
" the complete sbsenoe of the flrGt connecting convolation and tlin 
"uoncealment ofthe second, aa eBBBntiallycharBoteristic foatures in the 
" brain of thia animal, is by no means universally applicable. In only one 
" specimen did tha brain, in these particulars, follow the law which 
" Gratiolethas expressed. As regards the presence of the superior bridg- 
" ing convolution, X am inclined to think that it has existed in one hemi- 
" sphere, at ieaat, in a majority of the brains of thia animal which have, up 
■' tothis time, bten figured ordescribed. The superficial position of the 
" second bridging convolution is evidently lc«s frequent, and has as yet, 
" I believe, only been seen in the brain (A) recorded in this communis 
" cation. The asymmetrical arraugement in the ounvolotions of the 
'• two hemispheres, which pruviuus observers have referred to in their 
" liescciptions ia alaowellillastratedin these specimens." (pp. 8, 9.) 

Even were the preaenco of the temporo-oooipital, or eslemai per- 
pendicular, sulcus a murk of distinction between the higher apes and 
man, the vaiue of auch a distinctive character would be rendered very 
doubtful by the stmolure of the brain in the Platyrhine spes. In fact 
while the tempoio-oceipttal is one of the most constant of sulci in 
the Calarhiue, or Old World, apes, it ia never very strongly developed 
in the New World apes; it is absent in the smaller Platjrhini ; 
rudimentary in Pitkeeia ;" and more or leas obliterated by bridging 
convolutions in AUUs. 

A character which is thus variable within the limits of a single group 
can have no great taxonomio value. 

It IB further catabliBhed, that the d^reeofasymmetryof the convolu- 
tion of the two sides in tlie humiiu brain is aubjeof to much individual 
Tariation ; and that, in those individuEiJs of the Bushmaii race who hare 
been esamined, the gyri and anlci of the two hemispheres are consider- 
ably leas complicated and more symmetrical than in the European 
br^n, while, in same individuals of the chimpanzee, their complexity 
and asymmetry become notable. Thia is particularly tlie case in the 
brain of a young maJa chimpanzee figured by M. JBroca. (' L'ordre 
des Primates,' p. 165, fig. II.) 

Again, as respects the question of absolute size, it ia established that 
the difference between the largest and the smallest hcaitljy human 
brain is greater than the di£»«nce between the smallest healthy 
human brain and the largest chimpanzees or Clang's brain. 

MoreoveTj there is one circumstance in which the oiang's and chim- 
panzee's brains resemble man's, but in which they differ from the lower 
apes, and that is the prei«iite of two corpora candicantia — the 
CynumorpKa having but one. 

In view of these fa«fs I do not hesitate in thia year 1871, to repeat 
and insist upon the proposition which I enunciated in lg<]3." 

" So far as cerebral structure goes, therefore, it is clear that man 

" Kotus more especially on the " Flower 'On th« Anatomy i 

bridging convolutions la Ilie Brain Fithecia Monachas,' 'Proceedings i 

of ths Chimpanzee, 'Pi™eedings of the Zoological Society,' 1862. 

Uw Koyal Society of iidinturgti,' '' 'Man's Place in Nature,' p. 102 


303 The Descent of Man. Paut I 

■■ rtjffjrs less from the cLimpnuEfie ot the orang, than tlieae do bvhq 
" fioiii tho moDkeys, sod mat tlie difference between the brain of tha 
" cbimpaDiee nod otman is almost inaigniflcant, when uouiparud with 
" that between the chimpanzee brain and thnt of a Leniur." 

Ill the paper to which I hnye rtferred, Profesaor Biachoff docB not 
deny tlie lecond part of this etatement, but he first makes the irrelevant 
rcimarli: that it is not wonderful if the brains of an oraug and a Lemur 
BIO very different ; and secondly, goes on to assert that, " If we sucnes- 
- eively ooiapare the brain of a man witb that of bh orang: the biain of 
" this with that of a ohimpanzee ; of this with that of a frills, unii so 
"on of a ^jioiates, Sentnopittdctis, (^rnooepftoiuJi, Cefcopiifteem, Mocncus, 
" CWnis, (MUQtrin, Lemur, Sienops, flopute, we shall not meet with a 
"greater, or even as great a, break in (he degree of development of tho 
" criDvolnlions, as we find between tlie brain of a mau and that of an 
" "rang or chimpanzee." 

To which I reply, firstly, that whether this assertion be trae 
or false, it haa nutbiTjg wh»teTer to do with the proposition enuni^iuted 
in 'Man's Plaoe in Nature,' which refers riot to the development of tha 
convolutions alone, but to the stiuctnre of the whole brain. It Frofes»or 
Bisohoffhad taken the tronble to refer to p. 96 of the work he criticises, 
iu Eict, be would have found the following passage: "And it is a 
" remHrkalile eireomslanoe that thoogh, so far as our present knnw- 
" ledge extends, there is one tnio Btruolural break in tiie series of forma 
"of Simian brains, this hiatus does not lie between mau and the 
" manlike apes, but between the lower end the lowest Simians, or in 
"other words, between the Old and Now World apes and monkeys and 
"the Lemurs. Every Lemur whinh haa yet been examined, in fact, 
" has its cerebellum partially visible from above; and ita posterior lobe. 
" with the contained posterior cornu and hippocampus minor, wore or 
"less rudimentary. Every marmoset, Aiuerioan monkey, Old World 
" monkey, baboon, or manlike ape, on the contrary, has lis ecrebellum 
" entirely hidden, posteriorly, by the cerebral lobes, and possesses a 
" liuge posterior cornU with a weE-developed hippocampus minor." 

Thw statement was a strictly accurate account of what was known 
whan it was made; and it does not appear lo me to be more thau 
apparently weakened by the subsequent discovery of the relatively 
small development of the posterior lobes in the Siaman^ and in tha 
Howling monkey. Notwithstanding the exceptional brevity cjf tlio 
posterior lobes in these two species, no one will pretend that their 
brains, in the slightest degree, approach those of the I.emurs. And 
if, instead of putting SapuXe out of its natural place, as Professor 
liischoff most unaccountably does, we write the series of animals 
bo has chosen to mention as follows : Homo, PiYJieeus. TrogioA^im, 
Hylobales, SemnopUheom, Cynooepkalm, Cercopitkeea*, Macaout, Cebut, 
CallithriJh Uapale, Xemur, tilennpt, I venture to reaiSrm that the 
great break in this series lies between Hajude and Lemav, and that 
tills breaJt is considerably greater than that between any other two 
terms of that series. Professor Bischoff ignores the fact tlint Inn^ 
befoie he wrote, Gratiolet had Euggesti:d the separation of the Lei.iurs 
Irom the other FHrnatet on the very ground of the difference in their 
cerebral characters ; and that Professor Flower had made the followins; 
uboet^'utions in the course of his description of the brain of the Javaii 




Structure of tJie Brain. 


" And it ia espenialiy remaiiabie that, in the development of (ho 
^ poaterioi lobes, there JB no approximation to the Lemmiiie, short 
" hemiflphered, bmin, in those monkeyg which are oummonly supposed 
" to approacli this family in other respeota, ria., the lowet membora ol 
'■ the Platythine group, 

60 Sn as tie atracture of the adult brain ia coooerned, then, the very 
coofliderablo additions to our knowledge, which have been made by the 
researohes of bo many investigators, during tlie past ten yoata, folly 
justify the etatement which 1 made in 1^3, But it has t>een aa\ 
that, adioitting the similarity between the edult brains at iiUkU and I 
apes, they are nevectiieleBB, in reality, widely different, becanso they '• 
es:hibit fundamental difiorencea in the mode of their development. No 
one would be more ready than I to admit the force of flits argument, if 
euoh ftindamBntal differenceB of development really exist But I diny 
that thej do eiiat. On the contrary, there is a fundmnental agre&- 
ment in the deveJi^ment of the brain in men and apes. 

Gratiolet originated the atatement that there is a fundnmeutal 
difference in the developtnent of the bralna of apea and that of man — 
cODsiating r,\ thia; tha^ in the apes, the sulci which first make their 
appeorancti are situated on the posterior region of the cerebral hemi- 
Bpherea, while, in the human tcetuB, the Bulei first become visible on the 
mjiital lobes." 

This general statement is baaed upon two observationa, the one of a 
Gibbon almost ready to be born, in which the posterior gyri were " well 
"developed," while those of the frontal lobea were'- hardly indicated "" 
(]. c. p. S9), and the other of a human Itetua at the 22nd or 23nl week 
of uterogeatation. in which Qratiolet notes that the inaula was un- 
covered, but that noverthelees " dea incianiea semeut le lobe anterieuf, 
" une scisBUje pen profonde indique la separation dn lobe occipital, tree- 

"' "Chez tousles Binges, les plis 

Rolando, and one of the fronlitl 

" pi;st^rieurs se diveloppent les pre- 

" miers ; lea plis antfrieura se 

M. Alii, in his 'Notice eur les 

" vertebra occipitsle et la |>ari«tala 

ti'avaux anthropol<^iqueB de Gratio- 
let ■ QWa. de la Social d'Aathi-c 

"sont-elles relativement trfa-graudes 

pologie de Pari.,' 1868, page 32.), 

■' chc! le fiEtua. L'Homme prSsante 

writes thus : " Gratiolet a ea outre 

" les mains le cerrean d'un fcetis d( 

" i I'^poquo de I'apparitloa des plis 

" frontaai, qui soot les premiers 

" ptrienr, et telleraent rapprooh^ de 
" i'arang, que des naturaiistes tr^ 

" indiquis ; mals le develojipenient 

" g^n^ral du lobe frontal, envis^fi 

" saolament par rapport i sod 

" anthropoidas. M. Hmley, par ei- 

" Tolume.auit fesmemeslois qnedans 

" emple, n'hraite pas sur ce point. 

"les singes:" Gratiolet. >M^oire 

" Eh bien. c'est anr la eervean d'un 

eur les phs e£r£braux de rHoouue 

" fffitus de Gibbon qua Gratiolet a 

et des Primates,' p. 39, Tab. iv. 

fig. 3. 

" Giatiolet's words are (1. c. p. 

" iwsju'iin'eiistenipasMoor^ifeyfij 

39)! "Dans le f*tus dont il a'agit 

" sur is U^frwaui. 11 *tait don.; 

" bien auteria* i dira qua, d."l 

" bien diveloppia, taadis que les 

" plis dn lobe frontnl sont i peine 

" raiasent d'a en a, tandis que chei 

■' indiqnes." The figure, however 

"lea siDges elles se d^Telotpeiit 


204 ?"^ Descent of Man. Pabt T, 

" iHidiiit, d'niiieuis fics cette epoqne. Le ceate da la surBioe oerebrole 
" ent encore absDlui:ient lisse." 

Three views of this bmin are given in Plate II. figs, 1 , 2, 3. of ttie 
wnrk tited, shewing the upper, lateral and Inferior views of tbe henai- 
Bplieiea, but not the inner view. It ia worthy of note that the figure 
by no meims bears out Gralitlet's description, inasmuch aa tbe fissure 
(imtcrofcmpotal) oc tbe posterior half of the fhce of the bcmisphere. ia 
•BKiTS marked than any of those vaguely indicated in tlie anterior half. 
If tiie liguro is correct it in no way justifies Gratiolet's conolasion : 
" IlyttdonoentcBcescerveaux rthoseof [lOallithtiKand of a Gibbon] et 
" eel Di du fcalns humain une dimrence fnndamentoJ. Chez celui-oi, long- 
" temps avant qnn les plis temporaui apparaisaent, Jes piis fronteui 

tiinoe Gratiolet's lime, however, the development of the gyri and 
enlci of tbe brain has been made the subject of renewed investigation 
by Sclimidt, Biachoff, Pangch," and mora narticnlarly by Ecker," whose 
work is not only the latest, but by far the most romplete, memoir on 
ttie subject. 

The £nal resnlts of their inqniries may be summed op as follows ; — 

1. In the human foetus, the aylviao Hssuia is formed in the course of 
tlie third montb of uterogestation. In this, and in the fourth month, 
tlie cerebral hemiapheiea are smooth and rounded (with the exception 
of tbe sylvian depression), and they project liaekwards far beyond the 

2. The sulci, properly so called, begin to appear in the interval 
between the end of the fourth and the beginning of the sinth month of 
ffotal life, bat Ecber is carefal to point out that, not only the time, but 
the order, of t^eir appearance is subject to considerable individual 
variation. In no case, however, are either the frontal or the temporal 
sulci the earliest. 

The lirst which appears, in fact, lies on the inner face of tbe bemi- 
aphere (whenco doubtless Gratiolet, who lioes not seem to have examined 
that face in his fratus, overlooked it), and is either the internal pelpen- 
dicular (occipito-parietal). or the caii'arine sulcus, these two heitu; close 
together and eventually running into one another. Aa a rule the 
oocipito-parietal is the earlier of the two. 

3. At the latter part of this period, another sulcus, the " posterio, 
parietal," or " Fissure of Rolando " is developed, and it ia followed, in 
the course of the sisth month, by the other m-inoipal sulci of ihe 
fj-onta), parietal, tempoial and occipital lobes. There is, however, na 
clear evidence that one of these constantly appears before the otlier 
and it is remarkable that, in the brain at the period described and 
figured by Ecker (I. o. p. 212-13, Taf. D. figs. 1, 2, 3, 4), tbe antero- 
temporal sulcus (tci^iite 'pardSile) so cbaraoteristio of the ape's brain, 
IS as well, if not better developed than (he fissure of Kolando, and ia 
much mote marked than tbe proper frontal sulci. 

Taking the facts as they now stand, it appears to me thn! the order 
of the appearance of the sulci and gyri in the ftetai human brain is in 
perfect barmcny with the general doctrine of evolution, and with the 

' ' ZsT Eatwickelnngs Geschichte 

Furchen nnd Winlungcn der 

uroEsnirii-iiemisf._areu am drOjsiiira-Hemispharea im Fcetiu 

:hen nnd der Affen.' 'Archiv des Menachen.' ' Archiv filr Antbro- 

itr.ropolojrif,' iii., 1RS8. Hnipc,' Hi., 1888. 


CB«r. Vll. Structure of the Brain. SoJ 

view Uiftt man Iibs been evolved from Eome ape- like form ; tLougli there 
Can be do doubt that that form wus, in man; lespects, diffiuctit fiom 
buy laembei of the FrimaUt now living. 

Von Baer tanght us, half a century ago, that, in the course of theil 
devolopment, alfisd animals put on, at first, the characters of the greater 
groapa to which they belong, and, by degroea, DBaumetboaB which restrict 
them miain the limits of their family, genus, and epeciea ; and he 
proved, at the same time, that no devebpinentai stage of a higher 
animal is precisely simiiar b> the adult condition of aoy lower animal. 
It ia qiiil« correct to say that a frog poasea through the tondition of a 
fish, iaaamuch as at one period of its life the tadpole haa all the cha- 
racters of a fish, and, if it went no further, would have to be srouped 
anioDg fishes. But it ii9 equally true that a tadpole is very different 
irom any known fiEb. 

In like manner, the bruin of a hUTnac ftelus, at the fifth month, may 
correctly be said to be, not only the bmin of an ape, but that ol" an 
Arotopltheoine or marmoaet-like ape; for ita hemisphores, with their 
great posterior lobster, and with no sulci bat the sylvian and the 
(^Icarine, present Uie charaoteriatica found only in the group of the 
Arctopiliecine Prtmaiae. But it is equally true, aa Gmtiolet remarks, 
tliat,]n its widely open sylvian fiasure, it differs from the bisia of any 
actuid marmoset. No doubt it would be much more similar to the brain 
of an adviinccd fcetus of a mainioset. But we know nothing whatever 
of the development of the brain in the marmosets. In the Platyrhini 

K>per, tlie only observation with which I am acquainted ia due to 
nech, who found in the brain of a fcetal Cebua Apetla, in addition to 
the sylvian fissure aad the deep calcarine flBaure, only a very shallow 
atitoratemporjJ fissure (eciesiireparnHtte of Graliolet.) 

Now this fact, taken together with the circumstance that the antero- 
temporal sulcus is present in suoh Platyrhini as the Saimiri, which 
present mere traces of sulci on the anterior half of the estotior of the 
cerebral hemispheres, or none at all, undoubtedly, so far as it goes, 
affords fair evidence in favour of Gratiolet's hypothesis, that the 
posterior sulci appear before the ant«rLor, ia the brains of the 
FUityrhini. But, it by no means follows, that lie rule whidh may hold 
good for the Pwiythijii extends to the Gatarhini. We have no in- 
formation whatever respecting the development of the brain in the 
Cynnmor^ia; and, as r^ards the AnihTopmaorpha, noUiing but the 
aceouat of the brain of Uie Gibbon, near birth, already referred to. 
At the present moment, there is not a shadow of evidence to shew 
that the sulci of a chimpanzee's, or orang's, br^n do not appear in the 
same order as a man's. 

Gratiolet opens his preface with the aphorism. " II est dangereitx 
"dans les Bciences de conclure Irop vite." I fear he must have for- 
gotten this sound masim by the time he had reached the discussion of 
tha differences between men and apes, in the body of Ids work. Nc 
doubt, the exeelleril author of one of the most remarkable contributions 
to the just understanding of the mammalian brain which has ever been 
made, would have been the first to admit the insufScienoy of his data 
had he lived to profit by the advance of inquiry. The mistortnne is 
ihat Ma ooncluaiona have been employed by persons incompetent to 
appreciate their foundation, as arguments in favour of obscurantism." 

winiime et I'oriBin* de I'HoKmB 


206 The Descent of Man. fjtT t 

Bui it ia important to remark that, whether Gratioldt was right or 
wrong in Lis hypothesis respecting the relative order of appeaiaoRO of 
the lempoial aud frontal suloi, the fact retmabs ; that, before eithor 
tumporul or frontal sulci, appear, the fcetal brain of man presenla 
chamcterB which ore found only in the lowest group of the Pnmatei 
(leaving out the I-emurs) ; a.od that this ia exactly what we should 
eipeet to be the case, if Dian has resulted {rom the gradual modificH- 
tion of the Sams form at that from which the other FrimaUi hava 


t 207 ) 

Paet II. 



r^iKOiPLKS OF Sbxcal Sblectioh. 

nal characters — Sem 
if males— Polygamy— The m 
seinal Belection— Eagerness of the male— Vaiiahility of toe maie— 
Choice exerted by the female — Seiaal wmpared with uatnra) selection 
— InheritaDce, at corresponding periods of life, at corresponding seasoni 
of the yeai', and as limited by ses — Relations between the several forms 
of inheritance-— Causes why one £ex and the young are not modified 
through sexual selection — Supplement on the proportional numbers oi 
the two seies throughout the animal kingdom — The proportion of the 

■With animals which have their sesea separated, the males 
necessarO J differ from the females in their oi^ns of reproduction ; 
and these are the primary sesual characters. But the sexes 
often differ in what Hunter has called secondary sesnal cliarac- 
tors, which are not directly connected irith the act of reproduc- 
tion ; for instance, the male possesses certain organs of sense' or 
locomotion, of which the female m quite destitute, or' has them 
more highly-developed, in order that he may readily find or 
reach her ; or again the male has special organs of ptehenMon for 
holding her seciuely. These latter organs, of infinitely diTersified 
kinds, graduate into those which are commonly ranked aa 
primary, and in some cases can hardly he distinguished from 
thom ; we see instances of this in the complex appendages at thu 
apex of the ahdomen in male insects. Unless indeed we confine 
the term "primajy" to tie reproductive glands, it is scarcely 
possible to decide which ought to be called primary and whion 

The female often differs from the male in having organs for the 
nourishment or protection of hei young, such as the mupmarj 


2g3 Tile Descent of Man. PAat H 

glands of mammals, and the abdominal sacks of the marsupials. 
In some few cases also the male possesses similar organs, which 
are wanting in the female, such as the receptacles for the 
ova in certaia male fishes, and those temporarily developed, in 
certain male frc^. The females of most bees are provided with 
a special apparatus for collecting and caiTjing pollen, and theb 
ovipositor is modified into a sting for tlie defence of the larvie 
and the community. Many similar cases could be given, hut 
thoy do not here coocern us. There are, however, other sexual 
differences C[uite Tmconnected with the primary reproductive 
organs, and it with is these that we are more especially concerned 
— such as the greater size, strength, and pugnacity of the male, 
his weapons of offence or means of defence against rivals, his 
gau(ly colouring and various ornaments, his power of song, and 
other such characters. 

Besides the primary and secondary sexual differences, such aa 
the foregoing, the males and females of some animals differ in 
structures related to different habits of life, and not at all, or . 
only indirectly, to the reproductjve functions. Thus the females 
of certain flies (Culicidte and Tabanidie) are blood-suckers, 
whilst the males, Uving on flowers, have mouths destitute of 
mandibles.' The males of certain mollis and of some crustaceans 
((..;/. Tanais) have imperfect, closed months, and cannot feed. 
The complemental males of certain Cirripedes live like epiphytic 
plants either on the female or the hermaphrodite form, and are 
destitute of a mouth and of prehensile limbs. In these cases it ia 
the male which has been modified, and has lost certain important 
o^^ns, which the fcmalea possess. In other cases it is the female 
which has lost such parts ; for instance, the female glow-worm ia 
destitute of wings, as also are many female moths, some of which 
never leave theii cocoons. Many female parasitic crustaceans 
have lost their natatory legs. In some weevil-beetles (Curcu- 
lioiiidce) there is a great difference between the male and female 
in the length of the rostrum or snout ; " but the meaning of thia 
and of many analogous differences, is not at all understood. 
Difierences of strncture between the two sexes in relation to 
different habits of life are generally confined to the lower 
animals; but with some few birds the beak of the male differs 
from that of the female. In the Huia of New Zealand the 
difference is wonderfully great, and we hear from Dr. Bullcr 

' Westwood, 'Modorn Class, of " Kirby and Spence, ' lutiod no- 
Insects,' vol. ii. 1840, p. S4I. For tion In Enlomulsgy,' ToL iii. IB^e 
the i^tatemeot about Tanais, men- p. »0e. 
lioned below, I am indebt«i in Fjiti ' ' Birds of .\eitr Zealocd,' I S7J 


OnAi\ Vfll. Sexual Selection. 209 

that the male ases his strong beak in cliiselling the larva of 
■nseofe out of decayed wood, whilst the female probes the softei 
paita ■(rftli her iar longer, much cinved and pliant boak ; and 
thus thfiy njutually aid each other. In most cases, differences of 
structure between the eesas are more or less directly connected 
with the propagation of the epecies ; thus tt female, which has to 
nourish a multitude of ova, requires more food than the male, 
and consequently requires ejiecial means forpioouring it. A male 
animal, which lives for a very short time, might los« its oi^uB 
for procuring food through disuse, without detriment ; but he 
would retain his locomotive organs in a perfect state, so that 
ho might reach the female. The female, on the other hand, 
might safely loea her oj^ans for flying, swimming, or walking, 
if she gradually acquired habits which, rendered such powers 

CWe are, however, here concerned only with sexual selection. 
This depends on the advantage which certain individuals have 
over others of the same scs and species solely in respect of 
reproduction^ When, as in-the cases above mentioned, the two 
sesea differ in structure in relation to different habits of life, 
they have no doubt been modified through natural selection, and 
by inheritance limited to one and the game sex. So again tho 
primary sexual organs, and those fornourisbingor protecting the 
young, come under the same influence ; for those individuals which 
generated or nourished their offspring best, would leave, ojsteris 
•paribus, the greatest number to inherit their superiority ; whilst 
those which generated or nourished theii offspring badly, would 
kave but few to inherit their weaker powers. As the majobas to 
find the female, he requires organs of sense and locomotion, -ut 
if these organs are necessary for the other purposes of life, m in 
generally the case, they will have been developed through 
natural selection, "When the male has found the female, he 
sometimes absolutely requires prehensile organs to hold her; 
thus Dr. Wallace informs me that the males of certain moths 
cannot unite with the females if their tarsi or feet are broken. 
The males of many oceanic crustaceans, when adult, have their 
legs and anfennsJ modified in an extraordinary manner for the 
prehension of the female ; bonce we may suspect that it is be- 
cause these, animals are washed about by the waves of the 
open sea, that they require these organs in order to propagate 
their kind, and if so, their development has been the result of 
ordinary or natittal selection. Some animals extremely low in 
Ibe scale have been modified for this same purpose; tiius the 
males of certain parasitic worms, when fully grown, have the 
lower saIfao^ of the terminal part of tlieir bodies roughened 


The Descent of Man. 

like a rasp, and with thie they coil round and permanently hold 

When the two seses follow esactiy the same nabite of life, and 
the male has the sensory or locomotive oi^ans more higiily 
developed than those of the female, it may he that the perfeetiun 
of these is indispensable to the male for finding the female ; but 
in the vast majority of eases, they serve only to give one laalo 
an advantage over another, for with sufficient time, the less well- 
endowed males would succeed in pairing with the females ; and 
judging from the stracture of the femak, they would be in all 
other respects equally well adapted for their ordinary habits of 
life. Since in such cases the males have acquired their present 
structure, not from being better fitted to survive in the struggle 
for esistenoe, but from having gained an advant^e over other 
males, and from having transmitted this advantage to their male 
offspring alone, sexual selection must here have come into octioii. 
It was the importance of this distinction which led me to 
designate tiis fiinn of selection as SeiuaJ Selection. So again, 
if the chief service rendered to the male by his prehensile organs 
is to prevent the escape of the female before the arrival of other 
males, or when assaulted by them, these organs will have been 
perfected through semal selection, that is by the advantage 
acquired by certain individuals over their rivals. But in most 
eases of this kind it is impossible to distinguish between the 
effects of natural and sesual seleetaon. Whole chapters could 
be filled with details on the differences between the kcics in their 
sensory, locomotive, and prehensile organs. As, however, these 
structures are not more interesting than others adapted for the 
ordinary purposes of life I shall pass them over almost entirely, 
giving only a few instances under each class. 

There are many other structures and instincts which must 
have been developed through sesual selection —such as the 
weapons of offence and the means of defence of the males for 
fighting with and driving away their rivals— their courage and 
pugnacity— their various ornaments — their contrivances for pro- 

could not have been develo; 

8VUB ScJeatifiqae,' Feb, 1, 1873, 

65) B3 one fatal to the belief m through tha cnoice ot tne lemaiel 

lal selection, inasmmh a* lie Had I not met with this remark, I 

poses that I attribute all llie should not have thonght it poaeible 

:rence» between the saies to for any one to have read this chapMr 

isl selection. This distinguished and to have imagined that I main, 

iralist, therefore, like so many tain that the choice of the female 

tr Prenohmen, has not taken the had anything to do with the develop. 

ible Lo understand even the first mont of the prehensile organs In the 


Chap. YIII. Sexual Selection. 2 1 1 

ducii^ vocal or instrnmenlal muBic— and their glands for 
emitting odours, moat of these !att«r structnres serving only to 
allure or excito the female. It is clear that these characters are 
the reenlt of sexual and not of ordinary selection, since ■anarmed, 
unomamented, or unattractive males would succeed equally well 
in the battle fox life and in leaving a numerous progeny, but for 
the presence of better endowed males. We may infer that this 
would be the case, because the females, wijicb are unarmed and 
miomamented, are able to survive and procreate llieir kind. 
Secondary sesual characters of the kind just referred to, will bo 
fully discussed in the following chapters, as being in many 
respects interesting, but especially as depending on the will, 
choice, and rivalry 01' the individuals ol either sex. When we 
behold two males fighting for the possession of the female, or 
several male birds displaying their gorgeous plumage, and per- 
ibrming strange antics before an assembled body of females, wa 
cannot doubt that, though led by instinct, they know what they 
are about, and consciously exert their mental and bodily powers. 

Just ts man can improve the breed of his game-cocks by the 
selection of those birds which are victorious in the cockpit, so it 
appears that the strongest and moat vigorous males, or those 
provided with the best weapons, have prevailed under nature, 
and have led to the improvement of the natural breed or gpedes. 
A slight degree of variability leading to some advantage, how- 
ever slight, in reiterated deadly contests would aufflce for the 
work of sesual selection; and it is certain that secondary sexuaJ 
oharactera are eminently variable. Just as man can give beauty, 
accoi'ding to his atandajd of taste, to his male poultry, or more 
strictly can modify the beauty originally acquired by the parent 
species, can give to the Sebright banta,m a new and elegant 
plumage, an erect and peculiar carriage— so it appears that 
female birds in a state of nature, have by a long selection of the 
more attractive males, added fo their beauty or other attractive 
qualities. No doubt this implies powers of discrimiuatiou and 
taste on the part of the female which will at first appear 
extremely improbable; but by the facts to be adduced here- 
after, I hope te be able to shew Oiat the females actually 
have these powers. When, however, it is said that the lower 
animals have a sense of beauty, it must not be supposed that 
Buch sense is comparable with that of a cultivated man, with hw 
multiform and complex associated ide^. A more jiist eom- 
porison would be between the taste for the beautiful in animala, 
and that in the lowest savages, who admire and deck theinselvas 
with any brilUant, gbttering, or curious object. 

I'rom our ignorance on several points, the precise manner in 


212 The Descent of Man. Fart IL 

which, sexual selection acts is somowbat tuicertain. Neverthe- 
less if those naturalists who already helieve in the mntability of 
species, will read the following chapters, thoy will, I think, agree 
with me, that sexual selection has plajed an important part in 
the history of the organic world. It is certain that amongst 
almost all animals there is a struggle between the males for the 
possession of the female. This fact is so notorious that it would 
be Kuperfinons io give instances. Hence the females have the 
opportunity of selecting one out of several males, on the suppo- 
sition that their mental capacity suffices for the exertion of ii 
choice. In many cases special circumstanees tend to make the 
struggle between the males particularly severe. Thus the males 
of our migratory birds generally arrive at their places of breeding 
before the females, so that many males are ready to contend for 
each female. I am informed iQ" Mr. Jenner Weir, that tte bird- 
catchera assert that this is invariably the case with the nightin- 
gale and blackcap, and with respect to the latter he can himsulf 
confirm the statements 

Mr. Swajsland of Brighton has been in the habit, during the 
last forty years, of catching our migratory birds on their first 
arrival, and he has never known the females of any species to 
arrive before their males. During one spring he shot thirty-nine 
males of Bay's wagtail {Budytes Sail) before he saw a single 
female. Mr. Gould has ascertained by the dissection of those 
snipes which arrive the first in this country, that the males come 
before the females. And the like holds good with most of tlio 
migratory birds of the United States." The majority of the male 
salmon in onr rivers, on coming up from the sea, are ready to 
breed before the females. So it appears to be with frogs and 
toads. Throughout the great class of insects the males almost 
always are the lirst to emerge from the pupal state, so that they 
generally abound for a time before any females can be seen," 
The cause of this difference between the males and females ia 
their periods of arrival and maturity is eufflciently obvious. 
Those males which annually first migrated into any eoantry, or 
which in the spring were first ready to breed, or were the most 
eager, would leave the largest number of offspring ; and tliese 

' J. A- Allen, oa the 'Mammal* mdite plants are dichogamons ; that 

and WiatfT Birds of Florid^,' Boll, is, their male aad f«nials oi^hus ave 

«.*mp. Zoology, Hairard College, p. not ready at the same time, bo ths: 

268. they oaaflot be nelf-fertilifed. Now 

• Even with thnss plants in which in such flowers, the pollen is :r, 

the terns are ssparate, the male general matured before the stigma, 

flowers are generally mature be- thongh there are eiceptional ttises 

fore the temale. An first Ehen>a in wlilch the female organs are 

bj C. K. Spreagel, many hermaph- befiirehBDd. 


CHip. VIIl, Sexual SeUclion. 213 

wcjuld tend to inherit simiJar instincts acd constitutions. It 
must lie borne in mind that it would have been impossible to 
change very materially the time of eeiuft! maturity in the 
temales, without at the same time interfering with the period of 
the production of the youDg— a period which must be determined 
bj the seasons of the year. On the whole there can be no doubt 
that with almost all animals, in which the seses ore separate, 
there is a eonslanl3y recurrent stru^le between the males for 
the possession of the females. 

Our difficulty in regard to Bosual selection lies in undeiatand- 
ing how it is that the males which conquer other males, or thos9 
which prove the most attractiyo to the females, leave a greater 
nnrabar of offspring to inherit their superiority than their 
beaten and leas attractive rivals. Unless this result does follow, 
the characters which give to certain males an advantage over 
others, could not be perfected and augmented through serual 
selection. When the sexes esist in eiaetly equal numbers, the 
worst-endowed males will (except where polygamy prevails), 
ultimately find females, and leave as many offepring, as well 
fitted for their general habits of life, as the best-endowed males. 
Prom various fects and considerations, I formerly inferred that 
with most animals, in which secondary sesna! characters are 
well deTolox>ed, the males considerably exceeded the females in 
number; but this is not by any means always true. If the 
males were to the females as two to one, or as three to two, or 
oven in a somewhat lower ratio, the whole affair would be 
simple ; for the better-armed or more attractive males would 
leave the lai^st number of offspring. But after investigating, 
as far as possible, the numerical proportion of the seses, I do not 
believe that any great inequality iu number commonly exists. 
In most cases sexual selection appears to have been effective in 
the following manner. 

Let us take any species, a bird for instance, and divide the 
females inhabiting a district into two equal bodies, the one 
consisting of the more vigorous and better-nourished individuals, 
and the other of the less vigorous and healthy. The former, 
there can be httJe doubt, would be ready to breed in the spring 
before the others; and this is the opinion of Mr, Jenner Weir, 
who has carefully attended to the habits of birds during many 
years. (|rhere can also he no doubt that the most vigorous, 
best-nourished and earliest breeders would on an average 
succeed in rearing the largest number of fine offspring.' The 
males, a^ we have seen, are generally ready to breed before the 

' Heie is eicelleat eyidenee on an 'eiperienced oinithologiBt. Mr. 
tlu character of the oflfepring ft- ■ . ■" 


214 The Descent of Man. Pabt II 

femalos ; the strongest, and with Home species the Lest armed o( 
lie males, drive away tlie weaker ; and the former would then 
unite with the more Yigorous and better-nourished females, he- 
caiose they are the first to breed." Such yigorous pairs would 
Burely rear a larger niuaber of ofepring^tlian the retarded 
females, which, would be compelled to unite with the conqiiered 
and less powerful males, supposing the sexes to be numerically 
equal ; and this is all that is wanted to add, in the course of 
BuccessiTO generations, to the size, strength and courage of the 
males, or to improve their weapons.*^ 

But in very many cases the males ■which conquer their rivals, 
do not obtain possession of the femalea, independeatly of the 
choice of the latter. The courtship of animals is by no means 
80 simple and short an affeir as might be thought. The 
females are most escit«d by, or prefer pairing with, the more 
ornamented males, or those which are the best songsters, or play 
the best antics; but it is obviously probable that they would 
at the same time prefer the more vigorous and lively males, and 
this has in some cases been confirmed by actual observaticm.' 
Thus the more vigorous femalea, which are the first to breed, will 
have the choice of many males ; and though they may not always 
select the strongest or best armed, they will select those which 
are vigorous and well armed, and in other respects the most at- 
tractive. Both seses, therefore, of such early pairs would as above 
explained, have an advantage over others ia rearing ofEspriug ; and 
this apparently has sufficed during a long course of generations 
to add not only to the strength and fighting powere of the males, 
but likewise to their various ornaments or other attractions. 

In the converse and much rarer case of the males selecting 
particular females, it is plain that those which were the most 
vigorous and had conquered others, would have the freest 
choice ; and it is almost certain that they would select vigorous 
us well as attractive females. Such pairs would have an advaa- 

nd Winter Birds 

of E. Florida,' 

to those female bees whieh are the 

.229) of the later I, 

irooda, after the 

first to emerge from the pupa each 

cddentiLl destrnctii 

m of the tirst. 

yeai-. See bia remarkable esaay, 

ITS, thst these " a 

re found to be 

'AnwenduQg den Dai-win'sohen Lehre 

smaller and pulei 

aaf Bienen,' ' Verb. d. V. Jahrg.' 

those hatched earl 

« several broods 

• With respect to poultry, I havp 

rewiTed information, hereafter to 

tuie the birds of the earlier broods 

be given, to this effect, tiren with 

seem in all respect 

9 the most per- 

birds, such as pigeons, which pair 

feot and vigorous.' 

for life, the ffmnie, as 1 hear iroir. 

'Hermann Mdlle 

r has come to 

Mr. Jenner Weir, will desert bci 

his same conclusici 

IE with respect 

mate if he is injUFnd or gion-s weak. 


Chap. 7111. Sexual Selection. 215 

tagii in tearing offspring, more especially if the male had tUa 
I power to defend the female during the pairing-season ae occurs 
with some of the higher animals, or aided her in providing for 
the jotiLng. The same principles would applj if each ses pre- 
ferred and selected certain individuaJs of the opposite ses; 
supposing taat they Belected not only the more attractive, but 
likewiBe the mote vigorous individuals, 

Numerical Proportion cf the Two Sexes.— 1 have remarked that 
Kexual selection would be a simple affair if the males were con- 
siderably more numerous than tie females. Hence I was led to 
investigate, as far as I could, the proportions between the two 
seies of as many animals as possible; but the materials are 
Bcanty. I will here give only a brief abstract of the results, 
retaining the details for a supplementary discussion, so as not 
to interfere with the course of my argument. Domesticated 
animals alone afford the means of aacerbiining the propor- 
tional numbers at birth; but no records have been specially' 
kept fot this purpose. By indirect means, however, I have 
collected a considerable body of statistics, from which it appears 
that with most of our domestic animals the sexes are nearly 
equal at birth. Thus 25,560 births of race-horses have been 
recorded during twenty-one years, and the male births were 
to the female bhrths as 99-7 to 100. In greyhounds the in- 
equaUty is greater than with any other animal, for out of 6878 
births during twelve years, the male births were to the female 
as llO-l to 100. It is, however, in. some degree doubtfnl 
whether it is safe to infer that the proportion would be the same 
under natural conditions as under domestication ; for slight and 
unknown differences in the conditions affect the proportion of 
the sexes. Thus with mankind, the male births in Er^land 
are as 104'5, in Russia as 108'9, and with the Jews of Livonia as 
120, to 100 female births. But I shall recur to this curious point 
of the eicess of male births in the supplement to this chapter. At 
the Cape of Good Hope, however, male children of European 
extraction have been bom during several years in the proportion 
of between 90 and 99 to 100 female children. 

For our present purpose we are concerned with the proportion 
of the sexes, not only at birth, but also at maturity, and this 
adds another element of doubt; fur it is a well-ascertained fact 
that with man the number of males dying before or during birth, 
and during the first few years of infancy, is considerably larger 
than that of females. 80 it almost certainly is with male lambs, 
and probably with someother animals. The males of some species 
kill one another by fighttn; ; or they drive one another aboul 


2 1 5 The Desunl of Man. Part II. 

antil tkof become greatly emaciated. The? mufit ^!so bo often 
exposed to various dangers, whilst wanderii^ about in eager 
search for the females. In many kinds of fish the males are 
much smaller than the females, and tbej are beUeved often to be 
devonred by the latter, or by other fishes. The females of 
some birds appear to die earlier than the males ; thej arc 
also Uftble to be destroyed on their nests, or whilst in charge 
of their young. With insects tho female larvffi are often latter 
than those of the males, and would consequently be more likely 
to be devoured. In some cases the mature females are less 
active and less rapid in their movemente than the males, and 
could not escape so well &om danger. Hence, with animals in a 
state of nature, we must rely on mere estimation, in order to 
judge of the proportions of ijie sexes at maturity ; and this is 
bnt Uttle tniEtwortty, escept when the inequality is strongly 
marked. Nevertheless, as far as a judgment can be formed, we 
may conclude from the facts given in the supplement, that the 
'males of some few mammals, of many birds, of some fish and 
insects, are considerably more numerous than the females. 

The proportion between the sexes fluctuates slightly during 
successive yeaia: thus with race-horses, for every 100 mares bom 
the stallions variedfromlOT-l in one year to 926 in another year, 
and with greyhounds from 116-3 to 9d3. Bnt tad larger num- 
bers been tabulated throughout en area more extensive than 
England, these fluctuations would probably have disappeared; 
and Bucli as ttoy are, would hardly suffice to lead to effective 
sexual selection in a state of nature. Nevertheless, in the cases 
of some few wild animals, as shewn in the supplement, the 
proportions seem to fluctuate eitlier during different seasons 
or in different localitieB in a sufficient degree to lead to such 
selection. For it should be observed that any advantage, 
gained during certain years or in certain localities by those males 
which were able to conquer their rivals, or were the most 
attractive to the females, would probably be transmitted to the 
offspring, and would not subsequently be eliminated. During 
the succeeding seasons, when, from the equality of the sexes, 
every male was able to procure a female, the stronger or more at- 
tractive males prevjoualy produced would siill have at least as 
jiood a chance of leaving oflapring as the weaker or less attractive. 

Polygamy, — The practice of polygamy leads tothe same results 
as would follow fl;om an actual inequality in the number of the 
sexes ; for if eaeh male secures two or more females, many males 
cannot pair ; and the latter assuredly will be the weaker or less 
attractive individuals. Many mammals and some few birds nxi 


CMap. VIll, SeXttal Selsction. 2i; 

polygamous, but with aniniais belonging to the lower classes I 
have found ao evidence of this habit. The intelleetiial powers 
of Euch animals are, perhaps, not sufficient to lead them to 
collect and gaaid a harem of females. That some relation exists 
between polygamy and the development of secondary sexual 
oharaot<jrs, appears nearly certain ; and this supports the view 
that a numerical preponderance of males would be eminently 
favourable to the action of sexual selection, NeverflielesB many 
animals, which are strictly monogamous, especially birds, display 
(troDgly-marked BecoDdary sexual characters; whilst some few 
.inimals, which are polygamous, do not have such characters. 

We win first briefly run through the mammals, and then trnx 
to birds. The gorilla seems to be polygamous, and the male 
difiers considerably from (he female ; so it is with some baboons, 
which live in herds containing twice as many adult females as 
males. In South America tke Mycetes caraya presents well- 
marked eesnal diffierences, in colour, beard, and vocal organs; 
and the male generally lives with two or three wives : the male 
of the Cebua capudnus difiers somewhat from the female, and 
appears to be polygamous."' Little is known on this head with 
respect io most other monkeys, but some species are strictly 
monogamona. The ruminants are eminently polygamous, and 
they present sexual differences more frequently than almost any 
other group of mammals; this holds good, especially in their 
weapons, but also in other characters. Most deer, cattle, and 
sheep are polygamous ; as are most antelopes, though some are 
monogamous. Sir Andrew Smith, in speaking of the antelopes 
of South Africa, says that in herds of abont a dozen there was 
rateJy more than one mature male. The Asiatic AniU<^ saiga 
appears to ba the most inordinate polygamist in the world ; for 
Pallas" states that the male drives away all rivals, and collects a 
herd of about a hundred females and kids together ; the female 
is hornless and has softer hair, but does not otherwise differ 
much from the mate. The wild horse of the Falkland Islands and 
of the Western States of N. America is polygamons, but, except 
in his greater size and in the proportions of his body, differs but 
little from the mare. The wild boar presents well-marked sexual 

" On the Gorilla, Savage and Fasc. lii, 1777, p. 39. Sir Andrew 

IVymai 'Boston Journal of Hat. Smitli, ' Illnstratious of the Zoology 

Hist.' vol. V. 1845-47, p, 423. On of S. Africa,' I84B, pi. 29, on the 

Cjnoc8phalos,Brehm,'illnst.Thi«r- Kobaa, Owen, in his 'Anatomy of 

leben," B. L 1864, s. 77. On My- Vertebrates' (vol. iii. 1838, p. 833) 

eetes,Reiigger,'NatHreesch.!aaEe- gives a table shewing incidentally 

.1; D 1^ ioa/i_ J 14^ which species of antelopes are gre- 


ZIH The Descent of Man. 1'art IL 

cliaraetera, in his great tusks and some other points. In Europe 
and in India lie leads a solitajy life, eiMpt during the broeding- 
season ; but as is believed by Sir W. Elliot, who has had many 
opportunities in ladia of observing this animal, he conisorts at 
this season witii several females. Whether tJiis holds good 
m Europe is doubtful, bat it is supported by some evidence. 
The adult male Indian elephant, like the boar, passes much of 
his time in solitude ; but as Dr. Campbell states, when with 
others, " it is xiure to find more than one male with a whole herd 
" of females;" the larger males expelling or tilling the smaller 
and ircalcer ones. The male differs from the female in his immense 
tusks, greater size, strength, and endurance; so great is the 
difference in these respects, that the males when caught are 
valued at one-fiflh more than the females." The sexes of other 
pachydermatous animals differ very little or not at all, and, as 
far as known, they are not polygamists. Nor have I heard of any 
species in the Orders of Cheiroptera, Edentata, Insectivora and 
Eodents being polygamous, excepting that amongst the Eodcnts, 
the common rat, according to some rat-catchers, lives with several 
females. Nevertheless the two eesea of some sloths (Edentata) 
differ in the character and colour of certain patches of hair on 
their shoulders.'^ And many kinds of bats (Cheiroptera) present 
well-nmrked sexual differences, chieily in the males possessing 
odoriferous glands aod pouches, and by their being of a lighter 
colour." In the great order of Eodents, as far as I can learn, 
the sexes ra,relj differ, and when they do so, it is but slightly in 
the tint of the fur. 

As I hear from Sir Andrew Smith, the lion in South Africa 
sometimes lives witb a single female, but generally with more, 
and, in one case, was found with as many as five females ; so 
that he is polygamous. As far as I can discover, he is the only 
polygamist amongst all the terrestrial Carnivora, and he alone 
presents well-marked sesual characters. If, however, we turn 
to the marine Carnivora, as we shall hereafter see, the case is 
widely different; for many species of seals offer extraordinary 
sexual differences, and they are eminently polygamous. Thus, 
according to F6ron, the male sca^elephant of the Southern Ocean 
always possesses several females, and the sear-lion of Forster is 
said to be surrounded by from twenty to thirty females. In the 
North, the male sea-bear of Steller is accompanied by even a 

Pnw. Zoolog. Soc' 1873. 

" Dr. Camiibali, in ' Froc. Zco- 

" Dr. Gray, 

!oe.S«.'18Ga, p. 138, See also an 

MftB. of Nat. Hi) 

inierestlDg ixiiior, by Lieut. Job^i- 

'• See Dr. 

ilnne, in ' I'rot Asiati- Soe. of 

paper, in ' Pnw. 

Beagul,' May. \m%. 



Chap. "ViH. Sexual Selection. 219 

greater number of females. It ia an interesting foot, as Dr. 
Q-ill remarks,'* tliat in the monogamsus species, " or tboso 
" liTiDg in small communitieB, there is little difference in Ki?« 
" between the npJes and females ; in the social species, or ratlier 
" those of which the males have harems, the males are vastly 
" la]^er than the females." 

Amongst hirds, many species, the sexes of which differ greatly 
from each other, are certainly monogamous. In Great Britain 
we see well-marked sexual differences, for instance, in the wild- 
duek which pairs with a single female, the common blackbird, 
imd the bullfinch which is said to pair for life. 1 am informed 
by Mr. Wallace that the Uke is true of the Chatterers or 
CotingidsB of South America, and of many other birds. In several 
groups 1 have not been able to discover whether the species are 
polygamous or monc^amous.. Lesson says that birds of paradise, 
BO remarkable fortbeir6exuaidifferences,arepolygamous, but Mr. 
Wallace donbts whether he had sufficient evidence. Mi. Salvin 
tells me lie has been led to behevo that humming-birds are 
polygamous. The male widow-bird, remarkable for his caudal 
plumes, certainly seems to be a polygamist." I have been 
assured by Mr. Jenner Weir and bj others, that it is somewhat 
common for three starlmgs to frequent the same nest; hut 
whether this is a case of polygamy or polyandry has not been 

The GallinacesB exhibit almost as strongly marked sexual 
differences as birds of paradise or humming-birds, and many of 
the species are, as is well known, poljgamous ; others being 
strictly monogamous. What a contrast is presented between the 
sexes of the polygamous peacock or pheasant, and the mono- 
gamous guinea-fowl or partridge! Many similar cases could be 
given, as in the grouse tribe, in which the males of t!ie polj- 
gamons capercailzie and black-cock differ greatly from the 
females; whilst the sexes of the monogamous red grouse and 
ptarmigan differ very little. In the Cursores, except amongst 
the bustards, few species offer strongly-marked sexual dif- 
ferences, and the great bustard {Olu tarda) is said to be poly- 
gamous. With the Grallatores, extremely few species differ 
sexually, but the ruff (Madietes paynax) affords a marked 

'"The Eared Se-ls, ' Americaa Great Bustard, see L. Lloyd, 'Game 

Saturaliat,' vol. iv., Jan. 1871. Birds of Swedea," 1867, p. 19, aod 

" 'The Ibb,' vol. iii. 1861 p. 1S2. Mootngu and Scl by speak of 

palygamr of tiie Capetcailiia and 


220 The Dment of Man. pAii-r II. 

exception, and this species is bulieved by Montagu b> be a 
polygamist. Heaco it appears that amongst birds there often 
exists a close relation between polygamy and the development of 
strongly-marked sexuaJ differences. I asked Mr. Bartlett, of tho 
Zoological Gardens, who has had very large erperierice with 
birds, whether the male tragopan (one of the GaliinaceEe) waa 
polygamous, and I was struck by his answering, "I do not 
" know, but should think so firom his splendid colours." 

It deserves notice that the instinct of pairing with a single 
female is easily lost under domestication. The wild-duck is 
strictly monogamous, the domestic-duck highly polygamous. 
The Sev. W. D. Fox informs me that out of some lialf-tamed 
wild-ducks, on a large pond in his neighbourhood, so many 
mallards were shot by the gamekeeper that only one was left for 
every seven or eight females ; yet vmusually large broods were 
reared. The guinea-fowl is strictly monogamous ; but Mr. Fox 
finds that Lis birds succeed best when he keeps one cock to two 
or three hens. Canary-birds pair in a state of nature, but tho 
breeders in England successfully put one male to four or five 
females. I have noticed these cases, as rendering it probable 
that wild inon<^amons species might readily become either 
temporarily or permanently polygamous. 

Too little is known of the habits of reptiles and fishes toenable 
us to speak of their marriage arrangements. The 8tickIe-l>aok 
(Gasterosteus), however, is said to be a polygamist;" and tho 
male during the breeding season differs conspicuously from the 

To sum up on t!ie means through which, as far as wo can 
judge, sexual seleotion has led to the development of secondiiry 
sexual characters. It has been shewn that the largest number 
of vigorous oflspring will be reared from the pairing of tho 
strongest and best-armed males, victorious in contests over 
other males, with the most vigorous and best-nourished females, 
which are the first to breed in the spring. If such females select 
the more attractive, and at the same time vigorous males, they 
will rear a larger number of offepring than the retarded females, 
which must pair with the less vigorous and less attractive 
males. So it will be if the more vigorous males select tho more 
attractive and at the same time healthy and vigorous females; 
and this will especially hold good if the male defends the 
fijmale, and aids in providing food for the young. The ad- 
vantage thus gained by the more vigorous pairs in rearii^ a 
larger cumber of offepring has apparently sufficed to render 
sesuai seleotion efficient. But a large numerical preponderance 

" Nccl Huraplirejs, ' River Gardens,' 1857. 


Chap. Vm. 

Sexual Selection. 


of males over femalea will be still more efBoient; whether the 
preponderance is only occasional and local, or permanent; 
whether it occurs at birth, or afterwards from tlie greater de- 
struction of the females ; or whether it indireetlj follows from 
the practice of polygamy. 

The Male generally more Toodijied tluin the Female, — Throughout 
the animal Idi^dom, when the sexes differ in external appearance, 
it is, with rare exceptions, the male which Las been the more 
modified ; for, generally, the female retains a closer resemblance 
to the young of her own species, and to other adult members of 
the same group, prhe cause of this seems to lie in the males 
of almost all animals having stronger passions than the fomalos] 
Hence it is the males that fight together and sedulously display 
their charms before the females; aad the victors transmit their 
superiority to their male offspringj Why both sexes do not thus 
acquire tho characters of their fathers, will be considered here- 
after. That the males of all mammals eagerly pnrsue the 
females is notorious to every one. So it is with birds ; but laaiiy 
cock birds do not so much pursue fie hen, as display their 
plumage, perform strange antics, and pour forth their song in 
her presence. The male in the few iish observed seems much 
more eager than the female; and the same is true of alligators, 
and apparently of Batrachians. Ihroughoatthe enormous class oi 
insects, as Kirby remarks," "the law is, that the male shall seek 
" the female." Two good authorities, Mr. Blackwall and Mr. C. 
Spence Bate, tell me that the males of spiders and crustaceans 
are more active and more erratic in their habits than the females. 
When the organs of sense or locomotion are present in the one 
sex of insects and crustaceans and absent in the other, ^r when, 
fls is frequently the case, they are more highly developed in the 
one than in the other, it is, as far as I can discover, almost 
invariably the male which retains such ot^ans, or has tliem most 
developed ; and this shews that the male is the more active 
member in the eonrtship of the sexes." 

" Kirby and Spence, 'Introdno- 

females of this species are impreg- 

tion to EntomologJ,' vol. iii. 18'2fl, 

nated by the males which are born 

y. 342. 

in the same cells with them ; bot 

it is much more probable that the 

females visit other cells, so that 

Inwot*,' vol. ii. p. 160) forms »n 

eiception to the ml., ^ the male 

We shall hei'eafter meet in various 

haa rudimentary wings, and never 
qnits the cell in wh.cii it ia bom. 

classes, with a lew eiceptional ctbos, 

in which the female, instead of the 

irhlbt the female has well-developed 

male, is the scrker nnd wooer. 

wio)^ Audoain balieves that the 


232 TIte Desceitt of Man. Palit IL 

The female, on the other hand, with the rarest exceptions, is 
less eager than the male. As the illustrious Hunter ''° long a(jo 
obsened, she generally " requires to be courted ;" she is coy, and 
may oi'tea bo seen endeavouring for a long time to escape from 
the uinJe. Every observer of the habits of animaJs will be able 
to call to mind instances of this kind. It is shown by varioue 
facts, given hereafter, and by the results lairJy attributable to 
scinal selection, that the female, though comparatively passive, 
generally exerts some choice aad accepts one male in preference 
to others. Or she may accept, as appearances woiild sometimes 
lead U3 to believe, not the male which b the most attractive to 
her, but the one which is the least distasteful. The exertion of 
some choice on the part of the female seems a law almost as 
general as tho eagerness of the male. 

We are naturally led to enquire why the male, in so many ajid 
such distinct classes, has become more eager than tho female, so 
that he searches for her, and plays the more active part in court- 
ship. It would be no advantage and some loss of power if each 
sex searched for the other; hut why should the male almost 
always be the seeker? The ovules of plants after fertili- 
sation have to be nourished for a time; hence the pollen ia 
necessajily brought to the female organs — being placed on the 
stigma, by means of insects or the wind, or by the spontaneous 
movements of the stamens ; and in the AJgse, &c., by the loco- 
motive power of the SJitlierozooids. Witli lowly-oi^anised 
aquatic animals, permanently affixed to the same spot and having 
their scios separate, the male element is invariably brought to 
the female ; and of this we can see the reason, for even if the 
ova were detached before fertilisation, and did not require 
subsequent nourishment or protection, there would yet be greater 
difficulty in transportiEg them than the male element, because, 
being larger than the latter, they axe preduced in far smaller 
numbers. So that many of the lower animals are, in this I'e- 
e|)ect, analogous with plaats."' The males of affixed and aquatic 
animals havii^ been led to emit their fertilising element in 
this way, it is natural that any of their descendants, which 
rose in the scale and became locomotive, should retain the sarao 
liabit ; and they would approach the female as closely as pos- 
sible, in order not to risk the loss of the fertilising element in a 
long passage of it through the water. With some few of the lower 

I Knd female reptoJuclivc 


Chap. VIII. Sexual Selection. 223 

iinimals, tte females alone are fised, and tlie males of these must 
1)0 Hie seekers. But it is difficult to understand why tho males or 
Epeoies, of which the progenitors were primordially free, should 
invariably have acquired the habit of approaching the females, 
instead of being approached by them. But in al! cases, in order 
that the males should seek efHciently, it wonld be necessary that 
they should be endowed with strong passiens ; and the acquire- 
ment of such passions would naturally follow from the more 
eager leaying a larger number of offspring than the less eager. 

The great eagerness of the males has thus indirectly led to their 
much more frequently developing secondary sexual characters 
than the females. But the development of such characters 
would be much aided, if the males were more liable to varj than 
the females — as I concluded they were — after a long study of 
domesticated animals. Von Nathusius, who has had very wide 
esperienco, is strongly of the same opinion.^ Good evidence also 
in favour of this conclusion can be produced by a comparison 
of the two sexes in mankind. During the Novata Espedition'' 
a vast number of measurements was made of varioos parts of the 
body in different races, and the=men were found in almost every 
case to present a greater range of variation than the women; but I 
shall have to recur to this subject in a future chapter. Mr. J. 
"Wood," who has carefully attended to the variation of the muscle 
in man, puts in italics the conclusion that " the greatest number of 
*' abnormalities in each subject is found in the males." He had 
previously remarked that "altogether in 102 subjects, the varieties 
" of redmidancy were found to be half as many again as in 
" '—ales, contrasting widely wiih the greater frequency of 
ciency in females before described." Professor Macalister ' 
ise remarks " that variations in the muscles " are probably 
more common in males than females." Certain muscles which 
are not normally present in mankind are also more frequently 
developed in the male than in the female ses, although oscepiiona 
to this rule are said to occur. Dr. Bart Wilder ™ has tabulated 
the cases of 152 individuals with supernumerary digits, of which 
86 were males, and 39, or less than half, females, the remaining 
27 being of unknown sex. It should not, however, be overlooked 

" ' Vortrage Uber Viehiucht,' my ' Variition of Animals ami 

=>'R«ise der Novara: Anthro- 1868, r. 75. ' ' 

polog. Theil,' 1867, 9. 216-269. " ' Proteediags Royal Soc.' toI. 

The reeults were laloolflted ty Dr. ivi. Jul? 1868, jip. 519 and 534. 
Weisbach from measniements made " 'Proc Rojol liish Academy, 

by Di-s. K. Scherzer and Schwatz. vol. i. 1868, p. 123 
On the greater variability of the " ' Massaehnsotta Merlical Soc 

jiuias of domesticaleil animals, see vol, ii. No. 3, 186&, p, E. 


224 Tke Descent of Man. Part II. 

that women would more frequentlj emicavour ti> conceal a 
deformity of thiskind than men. Again, Dr. L.Meyer asserts that 
the ears of man are more variable in form than those of woman.^ 
Lastly the temperature is morevariablG Jnmiin than in woman.'' 

The cause of the greater general varialsility in the male sex, 
than in the female is unknown, except in so far as secondary 
sesual characters are extraordinarily variable, and are usually 
conSnod to the males ; and, as we shaJl presently see, thie fact is 
to a certain extent, intelligible. Throngh the action of sexual 
and natural selection male animals have been rendered ia very 
many instances widely different from their females; but in- 
dependently of selection the two sexes, from differii^ constitu- 
tionally, tend to vary ia a somewhat different manaer. Tlie 
female has to expend much oi^anic matter ia the formation of 
her ova, whereas the male expends much force in fierce coatcsts 
with his rivals, in wandering about ia search of the female, in 
), pouring out odoriferous secretions, &c. : and 

^ expenditure is generally concentrated within a short period. 
The great vigonr of the male diurii^ the season of love seems 
often to intensify his colours, independeatly of any marked dif- 
ference from the female.* In mankind, and even as low down 
in the organic scale as in the Lepidopfera, the temperature of the 
body is higher in the male than in the female, accompanied in the 
case of man by a slower pulse.* On the whole the expenditure 
of matter and force by the two sexes is probably nearly equal, 
though effected in very different ways and at different rates. 

From the causes just specified the two sexes can hardly fail to 
differ somewhat in constitution, at least during the breeding 
season; and, although they may ba subjected to exactly the 
same conditions, they will tend to vary in a different manaer. 
If such variations are of no Bervioe to either sex, they will not be 
accumulated and increased by soxual or natural selection. Never- 
theless, they may become permanent if the exciting cause act& 

Ant and nd t«iiticm by Uiem of the sperm- 

at Huid; bat this can hoi'dt; be 

ntl a h .jse : for manv male birds, for 

at bj- Dr. J. Stock n H ugh 

iD the 'Pop. Scien-e Re w j„., 

51, 1874, p. 97. " For mankind, see Dr. J. Stoct- 

'I'of. Mantegazza a 1 d i t n Hough, whose coneluBiona an, 

Tio per I'Aatbropologia,'' 1874, p. 97. Sea Gitard's obserT^ 

p. 306) that the bright tions on the Lepidoptera. as fs.y«i 

(, cotoroon in so man)' male ia tlie ' Zoological Record,' 1869, H 

I, are dae to the preseoca 347. 


OaAP. VIII. Sexual Selection. 225 

permanentlj ; and ia accotdinoj with a ftxqaent form of iiilipii- 
taiice tho7 may bo teansmitted to tliat sex alone in which they 
first appeared. In this case the two seses will come to preeeat 
permanent, yet imiinportant, differences of cliaracter. For 
instance, Mr. AUea shews that with a large nnmber of birds 
inhabiting the northern and eouthem United States, the speci- 
mens from the south are darker-coloured than those from the 
north ; and this seems to be the direct iiisiilt of the difierence in 
temperature, light, ijic, between the two regions. Now, in some 
few cases, the two sexes of the same species appear to have been 
differently affected ; in the Age!<Eiit phceniceus the males have had 
their colours greatly intensified in the south ; whereas with Car- 
dinaiie virginianus it is the females which have been thus affected ; 
with Quiscalus maj<n- the females have been rendered extremely 
variable in tint, whilst the males remain nearly uniform."- 

A few exceptional cases occur in Tarious classes of animals, in 
which the females instead of the males have acquired well 
pronounced secondary sexual characters, such as brighter colours, 
grea,ter size, strength, or pugnacity. With birds there has some- 
times been a complete transposition of the ordinary characters 
proper to each sex ; the females having become the more eager 
in courtship, the males remainir^ eompaiatively passive, but 
apparently selecting the more attractive females, as we may infer 
from the results. Certain hen birds have thus been rendered 
more highly coloured or otherwise ornamented, as well as more 
powerful and pngnacious than the cocks ; these characters being 
transmitted to the female offspring alone. 

It may be sv^ested that in some cases a double process of 
selection has been carried on; that the males have selected 
the more attractive females, and the latter the more attractive 
males. This process, however, though it might lead to the 
modiJioation of both sexes, would not make the one sei 
different from the other, imlesa indeed their tastes for the beauti- 
ful differed ; but this is a supposition too improbable io be worth 
considering in the case of any animal, excepting man. There 
are, however, many animals in which the seses resemble each 
other, both being fnmisiied with the same ornaments, which 
analogy would lead us to attribute to the agency of sesua] 
selection. In such cases it may be su^ested with more plausi- 
bility, that there has been a double or mutual process of sexual" 
selection; the more vigorous and precocious females selecting 
the more attractive and vigorous males, the latter rejecting all 
except the more attractive females. But from what we know 

" ' Mammali and Birda of E. Florida,* pp. 234, 380, 295, 


226 The Descent of Ma^i. Part II 

of tho habits of aBimala, this Tiew is Lardly probable, for tlie 
loale is generally eager to pair with any female. It is more 
probablfl that the ornaments eommon to both aexos were acc[iiirei' 
hj one sex, generally the male, and then transmitted to the oh- 
spring of both seses. If, indeed, during a lengthened period the 
males of any species were greatly to exceed the females in 
lumber, and then during another lengthened period, bat under 
different conditions, the reverse were to occur, a double, but 
not simultaneous, process of sexual selection might easily bo 
carried on, by which the two seses might be rendered widely 

Wo shall hereafter see that many animals exist, of which 
neither sex is brilliantly coloured or provided with special orna- 
ments, and yet the members of both seses or of taie alone ha^ve 
probably acquired simple colours, such as white or black, through 
sexual selection. Tho absence of bright tints or other ornaments 
may be the restdt of variations of the right kind never having 
occurred, or of the animals themselves having preferred plain 
black or white. Obscure tints have often been developed 
through natural selection for the sake of protection, and the 
acquirement through sexual selection of conspicuous colours, 
appears to have been sometimes checked fi'om the danger thuB 
incurred. But in other cases the males during long ages may 
have struggled together for the possession of the females, and 
yet no effect will have been produced, unless a larger number of 
offspring were left by the more successful males to inherit their 
superiority, than by the less successful : and this, as previously 
shewn, depends on many complex contingencies. 

Sexual selection acts in a less rigorous manner than natural 
selection. The latter produces its effects by the life or death at 
all ages of the more or less successful individuals. Death, indeed, 
not rarely ensues from the conflicts of rival males. But generally 
the less successful male merely fails to obtain a female, or obtains a 
retarded and less vigorous female later in the season, or, if poly- 
gamous, obtains fewer females ; so that the j leave fewer, less vigoiv 
ous, or no ofepring. In regard to structures acquired through 
ordinary or natural selection, there is in most cases, as long as the 
conditions of life remain the same, a limit to the amount of 
advantageous modification in relation io certain special purposes ; 
but in regard to structures adapted to make one male victorious 
over another, either in fighting or in charming the female, there 
is no definite limit to the amount of advantageous modification ; 
60 that as long as the proper variations arise the work of sexual 
selection wDl go on. This circumstance may partly account for 
the frequent and extraordinary amount of variabiUty presenteii 


Chap. VIII. Sexual Selection. 22/ 

by secondary sexual characters. Kevcrtlieless, natural selection 
■will determine tJiat Buct characters shall not be acquired by the 
victorious males, if tliey would bo highly injurious, either by 
expending too mnch of their vital powers, or by exposing them 
to any great danger. The development, however, of certain 
Btructures — of the horns, for instance, in certain stags— has been 
carried to a wonderful extreme ; and in some cases to im extreme 
which, as far as the general conditions of life are concerned, 
mast be sightly mjuriouH to the male. From this fact we leain 
that the advantages which favoured males derive from conquer- 
ing othec males in battle or courtship, and thus leaving a 
numerous progeiiy,are in the long run greater than those derived 
from rather more perfect adaptation to their conditions of life. 
We shall further see, and it could never have been anticipated, 
that the power to charm the female has sometimes been more 
important than tie power to conquer other males in battle. 


In order to understand how sexual selection has acted oa many 
animals of many classes, and in the course of ages has produced 
a conspicuous result, it is necessary to bear in mind the laws of 
inheritance, as far as they are known. Two distinct elements 
are inclnded under the term " inheritance "—the transmission, 
and tbe development of characters ; but as these generally go 
together, the distinction is often overlooked. We see this dis- 
tinction in those characters which are transmitted thro)^ 
the early years of Lfe, but are developed only at maturity 
or during old age. We see the same distinction more clearly 
with secondary sexual characters, for these are transmitted 
through both seses, though developed in one alone. That they 
are present in both sexes, is manifest when two species, having 
strongly-marked sexual characters, are crossed, for eaoh trans- 
mits the characters proper to its own male and female sex to tlw> 
hybrid offspring of either sex. The same fact is likewise mani- 
fest, when characters proper to the male are occasionally deve- 
loped in the female when she grows old or becomes diseaEcd, 
as, for instenee, when the common hen assumes the flowing tail- 
feathers, hackles, comb, spurs, voice, and even puguacify of the 
cock. Conversely, the same thing isevidont, more or less plainly, 
with castrated males. Again, mdependently of old age or disease, 
characters are occasionally transferred from the male to the 
female, as when, in certain breeds of the fowl, spurs regularly 
appear in the young and healthy females. But in truth they are 
pimply developed in tiic female; for in every breed eacb detail 


228 The Descent of Man. Part n. 

in the stiuctnre of the spur is transmitted throngli tlie ftraale 
to her male offspriDg. Macy cases wjil hereafter be given, where 
tte female exhibits, more or less perfectly, characters proper to 
the maJe, in whom they must hare been first developed, and then 
transferred to the female, Tha converse case of the first de- 
velopment of characters ia the female and of transference to tha 
male, is less frequent; it will therefore be wail tn give one strik- 
ing instance. With bees the pollen-collecting apparatus ia nsed 
by the female alone for gathering pollen for the larvie, yet in 
most of tha species it is partially developed in the males 
to whom it is quite useless, and it is perfectly developed 
in the males of Bombns or tha humbie-beo." As not a 
single other Hymenopi«rouE insect, not even the wasp, which is 
closely allied to the bee, is provided with a pollen-collecting 
api)arBtu8, we have bo grounds for supposing that male beea 
primordially collected pollen as well as the females ; although 
we have some reason to suspect that male mammals primordially 
suckled their young as well as the females. Lastly, in all cases of 
reversion, chajacters are transmitted through two, three, or many 
mora generations, and are then developed under certain unknown 
favourable conditions. This important distinction between 
transmission and development will be best kept in mind by the 
aid of the hypothesis of pangenesis. According to this hypothesis, 
every unit 01 ceU. of the body throws offgemmulDSoriiiideveloped 
atoms, which are transmitted to tha ofispring of both sexes, and 
arc multipUed by self-division. They may remain ondeveloped 
daring the early years of lite or during successive generations; 
and their development into units or cells, like those from which 
they were derived, depends on their affinity for, and union 
with other units or cells previously developed in the due order 
of growth. 

Inheritance at correipondinij Periods of Zi/e.—lhis tendency 
is well established, A new character, appearing in a yoimg 
animal, whether it lasts thioughont life or is only transient, will, 
in general, reappear in the o3?pring at the same age and last 
for the same time. If, on the other hand, a new character 
appears at maturity, or even during old age, it tends to re- 
appear in tho offspring at the same advanced age. When devia- 
tions from this rule occur, the transmitted characters mucli 
oftensr appear before, thau after the corresponding age. As I 
have dwelt on this subject sufficiently in another work,'' I will 

" H. Mililep, 'AnwenduDg der "• -The Variation of Animals 

Danrin'echeli Lehre,' lie. Verh, and Plants nndcp Domestication,' 
i. n. V. Jahrg uii. p. 43. voL ii. 1868, p. 75. In th» lait 


Chap. VJIT. Sexual Selection. 229 

here meroJj give two or three jostancea, for the sake of recalling 
the subject to the reader's mind. In several breeds of the Fowl, 
the down-covered chickens, the young birds in tliett first true 
plunu^e, and the adults differ greatly from one another, as well 
as from their common parent-fanu, the GoUks bankiaa; and 
these characters are feithfully transmitted by each hroeii to their 
offspru^ at the corresponding periods of life. For instance, the 
chickens of spangled Hambui^, whilst covered with down, have 
a few dark spots on the head and rump, but are cot striped 
longitudinally, as in many other breeds ; in their first true plu- 
mage, " they are beautifully pencilled," that is each feather is 
transversely marked by mimerous dark bars; but in their second 
plumage the feathers all become spangled or tipped with a dark 
round spot'* Hence in this breed Tsiiatioae have occurred at, 
and been transmitted to, three distinct periods of life. The 
Pigeon offers a more remarkable case, becaiise the aboriginal 
parent species does not undergo any chai^ of plumage with 
iidvancing age, excepting that at maturity the breast becomes 
more iridescent ; yet there are breeds which do not acquire their 
characteristic colours until they have moulted two, three, or 
four times; and these modifications of plumage are regularly 

Inheritance at corresponding Seasons of the Tear. — With animals 
in a state of nature, innumerable instances occur of characters 
appearing periodically at different seasons. We see this in the 
horns of the stag, and in the fur of arctic animals which becomes 
thick and white durii^ the winter. Many birds acquire br^ht 
coloiffls and other decorations dni'ing the breeding-season alone. 
Palias states,*" that in Siberia domestic cattle and horses become 
lighter-coloured during the winter ; and I have myself observed, 
and heard of similar strongly marked changes of colour, that is, 
from brownish cream-colour or reddish-brown Ut a perfect white, 
in several ponies in England. Although I do not know that this 
tendency fo change the colour of the coat during different season 3 

chapter bat one, tbs proTisioDal 

hypothesis of paDgfmesis, above toL ii, p. 77. 

alluded to, is fully explained. " ' Novie apeciea Quadrupedoi 

" Tbase facts are givea oq the Glirium oriiifle,' 1778, p. 7. 

high authority of a great breeder, the transmission of oolonr by 

Mr.TeebaviseeTegetmeier's'Ponl- h^-rse, see 'Variation of Anim 

try Book,' 1868, p. 158. On tJie Sic, under Domestication,' voL i. 

chai'acters of chickens of different 51. Also vol. ii. p. 71, for a gi 

breeds, and on the breeds of the ral disonssion on ' Inheritance 

pigeon, alluded to in the following limited by Sex." 
pirngmph, see 'V.irialion of Ani- 


230 me Vesunt jj Mafi. PiuT IL 

IS traiismitted, jet it proliably is so, as all shades of coloiW aie 
stronglj interited by tte horse. Nor is this form of inheritaiico, 
as iimited bj the seasons, more remarkable than its limitation 
by age or sex. 

InheriUaice as Limited ly Sex. — The eqnal tranamissioil ol 
chajacters ia both seses is the commonest form of inheritance, 
at least with those animals which do not present etrongly-marked 
sexual differences, and indeed with many of those. But characters 
are somewhat commonly transferred exclusiyely to that sex, ia 
■whici they first appear. Ample evidence on this head has been 
advanced in my work on ' Variation under Domestication,' but a 
few instances may here be given. There are breeds of the sheep 
and goat, in, whidi the horns of the male differ greatly in shapo 
from those of the female ; and these differences, acquired under 
domestication, are regularly transmitted to the same sex. As a 
rule, it ia the females alone in cats which are tortoise-ahell, 
the corresponding colour in the males being rusty-red. With 
most breeds of the fowl, the characters proper to each ses 
are transmitted to the same sex alone. So general is this form 
of transmission that it is an anomaly when variations im certain 
breeds are transmitted equally to both sexes. There are also 
certain sub-breeds of the fowl in which the males can hardly be 
distinguished from one another, whilst the females differ con- 
siderably in colour. The sexes of the pigeon in the pajen1>-specie8 
do not differ in any external character; nevertheless, in certain 
domesticated breeds the male is colonied differently from tho 
female.'* The wattle in the English Carrier pigeon, and the crop 
in the Pouter, are more highly developed in the male than in the 
female ; and although these rfiaracters have been gained through 
long- continued selection by man, the slight differences between 
the sexes are wholly due to the form of inheritance which has 
prevailed ; for they have arisen, not from, but rather in opposi- 
tion to, the wish of the breeder. 

Most of OUT domestic races have been formed by the accumula- 
tion of many slight variations ; and as some of tho successive 
steps have been transmitted to one sex alone, and some to both 
sexes, we find in the different breeds of the same species all 
gradations between great sexual dissimilarity and complete 
similarity. Instances have already been given with the breeds 
of the fowl and pigeon, and under nature analogous cases aie 

" Dr. Chapois, ' Le Plgson Voj-a- . similar diffeifncos in certBin brccelj 
g«nr Beige,' 18K5, p. 87. Boitard at McdeD», ' he yariaziooi dei 
et Corbie, ' Lea Pigeons de Vobira,' Colombi domestici,' del Piolo Bo- 
fa., 1824, p. 173. See, also, on niz:!!, 1873. 


Chap. Till. Sexual Selection 23! 

With animals nitder domestication, but whether in 
nature I will not Tonture to say, one sox may lose charactera 
proper to it, and may thus come somewhat to resemble tho 
opposite sei ; for instance, the males of some breeds of the fowl 
have lost their masctiline tail-plumes and tackles. On the 
other hand, the differences between the sexes may be increased 
under domestication, as with merino sheep, in which tho 
ewes have lost their boms. Again, cbai'acters proper to one 
sei may suddenly appear in the other sex ; as in those sub- 
breeds of the fowl in which the hens acquire spurs whilst jonng ; 
or, as in certain Polish sub-breeds, in which the females, as 
there is reason to believe, originally acquired a crest, and sub- 
bequently transferred it to the males. All these cases are iu- 
t«lligibl6 on the hypothesis of pangenesis ; for they depend on 
the gemmulea of certain parts, although i 
becomii^, through the ii " 
or developed in either sex. 

There is one diflcult question which it will be convenient to 
defer to a future chapter; namely, whether a character at first 
developed in botli sexes, could through selection be limited in 
its development to one sex alone. If, for instance, a breeder 
observed that some of his pigeons (of which the characters are 
usually transferred in an equal degree to both sexes) varied into 
pale blue, could he by loag-continned selection mate a breed, 
in which the males alone should be of this tint, whilst the females 
remained imchanged? I will here only say, that this, though 
perhaps not impossible, would be extremely difficult; for tbe 
natural result of breeding from the pale>blue males would be 
to change the whole stock of both sexes to this tint. If, how- 
ever, variations of the desired tint appeared, which were from 
the first limited in their development to the male sex, there would 
not be the least difficulty in making a breed with the two sexea 
of a different colour, as indeed has been effected with a Belgian 
breed, in which the males alone are streaked with black. In a 
similar manner, if any variation appeared in a female pigeon, 
which was from the first sexually limited in its development to 
the females, it would be easy to make a breed with the females 
alone thus characterised ; but if the variation was not thos 
originally limited, the process would be extremely difficult, per- 
haps impossible." 

" Since the publication of the perieneed a breeder hb Wr. Teget- 
fii,-.t edition of this work, it has meier. After doscribing some cu- 
t^en highly satisfactory to me t« 
6aJ the following vcmaiks (the 

■ts m pigeons, of thf 

■ trn 





232 The Deueut of Man. I'iet II 

On ihi Relalion between, the Pm-iod i^ Devdopmenl of a Character 
ahd its Transmission to oae Sex or to both Sexes. — WLy certain 
characters stould be inherited by both seses, and other charac- 
ters by one ees alone, namely by that ses in wliicli tlie character 
first appeared, is in most cases quite imlmown. We cannot even 
conjectnre why with certain sub-breeds of the pigeon, black 
BtrJEB, though transmitted through the female, should be deve- 
loped in the male alone, whilst every other character is equally 
transferred to both seses. Why, again, with cats, the tortoise- 
shell colour should, with rare eseeptions, be developed in the 
female alone. The very same character, such ss deficient or su- 
pernumerary digits, colour-blindness, &a., may with mankind bo 
inherited by the males alone of one family, and in another family 
by the females alone, though in both cases transmitted through 
the opposite as well as through the same ses.^' Although we aro 
thus ignorant, tte two following mice seem often to hold good— 
that variations which first apjje-aJ- in either sex at a late period of 
life, f«nd to be developed in the saine sexalone; whilst variations 
which first appear early in life in eitiaor ses tend to be developed in 
both sexes. I am, however, far from supposing that this is the 
sole determining cause. As I have not elsewhere discussed this 
subject, and as it has an important bearing on sexual seiectiou, 
I must here enter into lengthy and somewhat intricate details. 

It is in itself probable that any character appearing at an 
early ago would tend to be inherited equally by both seses, for 
the seses do not differ much in constitution before the power 
of reproduction is gained. On the other hand, after this power 
has been gained and the sexes have come to differ in constitution, 
the gemmules (if I may again use the language of pangenesis) 
which aro east off from each varying part in the one ses would 
lie much more likely to possess the proper affinities for imitjng 
with the tissues of the same ees, and thus becoming developed, 
than with those of the opposite sex. 

I was first led to infer that a relation of this kind exists, from 
the fact that whenever and in whatever manner the adult malo 
differs from the adult female, he differs in tJie same manner from 
the young of botli seses. The generality of this fact is quite 
remarkable: it holds good with almost all mammals, birds. 

cith this character, ho siijs : "It ix 

" facts that I have related ; but it 

' a smgukr circumstance that Mr. 

" U remaikable how vwy olosely 

' Darwia should hnve suggested the 

" he suggested the right icethod ot 

' poasibililj of modifyiQg the seiuaj 

■ colours cf birds ij » ooorse of 

' aitiBcial selection. When he did 

Variation of Animals uodel' Dome-- 

' >D, be Vis in ignorauc; of these 

tiiation,' vol. ii. p. 72. 


Chap. Vm Sexual Sekciion. 233 

amphibians, and fishes ; also witii many crasfaceans, spiders, and 
some few insects, sucli as certain orthoptera and libellnlEs. In 
all these cases the variaSions, through the accumulation of which 
the male acquired his proper masculine characters, must have 
oocrared at a somewhat lata period of life ; otherwise the young 
males would have been similarly characterised ; and conformably 
with oni rule, the Tariations are transmitted to and developed in 
the adult males alone. When, on the other hand, the adult male 
closely resembles the young of both sexes (these, with rare 
exceptions, being alike), he generally resembles the adult female; 
and in most of these cases the variations through which the yonng 
and old acquired their present characters, probably occurred, 
according to oar rule, during youth. But there ia here room for 
doubt, fur characters are sometimes transferred to the ofepring 
at an eaiher age than that at whieh they first appeared in the 
parents, so that the parents may have varied when adult, and 
have transferred their characters to their offspring whilst young. 
There are, moreover, many animals, in which the two sexes closely 
resemble each other, and yet both differ irom their young ; and 
here the character of the adiills must have been acquired late in 
life ; nevertheless, these characters, in apparent contradiction 10 
our rule, aro transferred to both sexes. We must not, however, 
overlook the possibiUty or even probabiUty of anccessiva varia- 
tions of the same nature oociu;ring, under exposure to similar 
conditions, simultaneously in both sexes at a rather late pieriod 
of life ; and in this cose the variations would be transferred to 
the oHapring of both seses at a corresponding late age ; and there 
would then be no real contradiction to the rule that variations 
occurring late in life ate transferred exclusively to the ses in 
which they first appeared. This latter rule seems to hold true 
more generally than the second one, namely, that variations 
which occur in either sex early in hfe tend to be transferred to 
both sexes. As it was obviously impossible even to estimate in 
how large a number of cases throughout the aniinal kingdom 
these two propositions held good, it occurred to me to investigate 
some striking or crucial instances, and to rely on the result. 

An excellent case for investigation is afforded by the Deer 
family. In all the species, but one, the horns are developed 
only in the males, though certamly transmitted through the 
females, and capable of abnormal development in tliem. In the 
reindeer, on the other hand, the female is provided with horns ; 
so that in this species, the horns ought, according to our rule, 
to appear early in life, loi^ before the two sexes are mature 
and have uome to differ mtich in constitution. In all the 
ulhor species the horns o-jght to appear later in lifis, which 


234 1'^ Descent of Man. Pabt IL 

would lead to their deyelopmcnt in that sex alone, in which 
tiiey first appeared In the progenitor of the whole Family. Now 
in seven species, helongicg to distinct sections of the family and 
inhabiting difierert regions, ia which the stags alone hear horns, 
I find that the horns first appear at periods, varying from nine 
months after birth in the roebuck, to ten, twelve or even more 
months in the stags of the sis other and larger species." But 
with, the reindeer the case is widely different ; for, as I hear from 
Prof. Nilsson, who kindly made special enqniries for me in 
Lapland, tiie horns appear in the young animals within four or 
five weeks after birth, and at the same time in both seies. So 
that here we have a structure, developed at a most unnsually 
early age in one species of the family, and litewiso common to 
both sexes in this one species alone. 

In several kinds of antelopes, only the males are provided with 
horns, whilst in the greater number both sexes bear horns. 
With respect to the period of development, Mr. Blyth informs 
me that there was at one time in the Zoological Gardens a young 
koodoo (jin(. sirepsjceros), of which the males alone are homed, 
and also the young of 'a closely-allied species, the eland {^Ant. 
arms), in which both sexes are horned. Now it is in strict 
conformity with our rule, that in the young male koodoo, 
although ten months old, the horns wore remarkably small, con- 
sidering the size ultimately attained by them; whilst in the 
young male eland, although only three months old, the horns 
were already very much lai^r than in the koodoo. It is 
also a noticeable fact that in the prong-homed antelope," 
only a few of the females, about one in five, have horns, and 
these are in a rudimentary state, though sometimes above four 
inches long; so that as far as concerns the possession of horns 
by the males alone, this species is in an intermediate condition, 
and the horns do not appear until about five or sis months after 
birth. Therefore in comparison with what little we know of 
the development of the horns in other antelopes, and from what 

» I am mach obliged to Mr. tinent, see J. D, Caton, in 'Ottawa 
Cnpples for having made enqniries Acad, of Kat. Se. 1868, p. 13. Kor 
for me in regard to the Roebuck Cer^va Eldi of Pegu, see Lieut, 
tnd Red Deer of Scotland from Mr. Beavan, 'Proe. Zoolog. Sea.' IBBT, 
Robertson, Clie eiperienced head- p. 762. 

forester to the Marquis of Breadal- " Antilocapra Americana. I have 

bano. In regard to Fallow-deer, I to thank Dr. CanSeld for informa- 

others for information. For the femde: see aino his paper in 'Proc. 

Cs-WM alces of N. America, see Zoolog. Soc' 1866, p. 1C9. Alsc 

■lam) and Water,' 1888, pp. 221 Owen, 'Anatomy of Vertebrate*, 

«ad 2S* ( and for the C. Virgini-Bma vol. iii. p. B27. 
iDa iinatgylooeros of the same eoo- 


CHAr. Vin. Sexttal Selection. 235 

we do know with respect to the horns of deer, cattle, <to., tlioio 
of the prous-iomed antelope appear at an intermediate period 
of life,— that is, not very early, as in cattle and sheep, nor very 
late, as in the larger deer and antelopes. The horns of sheep, 
goats, and cattle, which are well developed in both sexes, though 
not quite equal in size, can be felt, or even seen, at birth or soon 
afterwards." Our riHe, however, seems to fail in some breeds 
of sheep, for instance merinos, in which the rams alone are 
horned; for I cannot find on enqniry," that the horns are 
developed later iu Ufb in this breed than in ordinary sheep in 
which both seses are horned. But with domesticated sheep the 
presence or absence of horns is not a firmly fised character ; for 
a certain proportion of the merino ewes bear small horns, and 
eome of the rams are hornless; and in most breeds hornless 
ewes are occasionally produced. 

Dr. W. Marshall has lately made a special stndy of the pro- 
tuberances so common on the heads of birds," and he comes 
to the following conclusion ;— that with those sjiecies in which 
they are confined to the males, they ate developed late in 
life; whereas with those species ia which they are common to 
the two seses, they are developed at a very early period. This is 
certainly a striking confirmation of my two laws of inheritance. 

In most of the species of the splendid family of the Pheasants, 
the males differ conspicuonsly from the females, and they acquire 
their ornaments at a rather late period of life. The eared 
pheasant (CroBsop's^ aaritaiit), however, offers a remai-kable 
exception, for both seses possess the fine caudal plumes, the 
large ear-tufts and the crimson velvet about the head ; I find 
that all these characters appear very early in life in accordance 
with rule. The adult male can, however, bo distinguished from 
the adult female by the presence of spurs; and conformably 

" I have heeo assured that the 

however, a breed of sheep in which, 

horns of the sheer i" North Wales 

as with merinos, the cams alone 

tan always b* felt, and are some- 

bear horns; and Mr. WinwcoJ 

times even an inch in length, at 

Reade informs toe that in one casu 

birth. Yonatt says ('CatUe,' 1834, 

obBei-ved by him, a young ram, 
born ou Feb. 10th, first eiiewe.t 

the frontal bone in cattle penetr.if*s 

horns on March Sth, so '.hat in this 

tho cutis at birth, and that the 

instance, in conformity with rule. 

horny matter b soon formed over 

the development of the horns oc- 


curred at a later period of life thaa 

" I am greatly indebted to Prof. 

in Welsh Bheep, in which both seies 

Victor Cams for hating made en- 

are horned. 

<)uiii85 for me, from the highest 

" 'UeberdicknBcheTnenSchadel- 

aothoFities, with respect to the 

hocker der Viige! ' in the ' Nieder- 

merino sheep of Saiony, On the 

landiscneB Archiv fill' Zooiogie,' 

Sninea coast of Africa there ie. 

Bandl, Heft3, 1872. 


The Descent of Man. 

Paet II 


with our rule, these do not begm to he developed before the ago 
of sis months, as I am assured hj Mr. Bartlott, and e¥en at this 
age, the two seses can liardlj be distiDgnished." The male and 
female Peacock differ conspicnouBly from each other in almost 
every part of their plumage, escept in the elegant head-crest, 
which is common to both seses; and this is developed very earlj- 
in life, long before the other ornaments, which areeontined tothe 
male. The wiid-duck offers an anali^ons case, for the beautiful 
green speculnm on the wings ib common to both seses, though 
duller and somewhat smaller in the fcmaJo, and it is developed 
early in life, whilst the curled tail-feathers and other ornaments 
of tie male are developed later." Between such extreme cases 
of close sexual resemblance and wide dissimilarity, as those of 
the Crossoptilon and peacock, many intermediate ones could be 
given, in which the characfcrs follow our two rules in their order 
of developments 

As most insects emerge from the pupa! state in a mature 
condition, it is doubtful whether the period of derelopment can 
determine the transference of their characters to one or to both 
Bescs. But we do not know that the coloui'ed scales, for instance, 
in two species of butterflies, in one of which the sexes differ in 
colour, whilst in the other they are alike, are developed at the 
eame relative age in the cocoon. Nor do we know whether all 
the scales are simultaneously developed on the wings of the same 
species of butterfly, in which certain coloured marks are confined 

" la the common peaoMk (Paoo 

seies; bat I hare not been able to 

eratatag) tlie male alone possesses 

distoier whether ita fnll develop- 

spurs whilot both seies of the Java 

manl occurs later in life in the 

Pe.iWK;k iP mutioBs) offer the ujj 

males of such species, than in the 

u ual case of being lurnieherl with 

mde of the common duet, as ought 

.pais Hence I fnlly eipected tl at 

to be the case according to our 

in the latter tpecies they would 

rule With the allied Mergva c«- 

h-ite been developed earber m life 

cdUi^t we have, however, a case of 

than in the common peacock bnt 

this kind r the two seies differ con- 

III Hegt of Amsterdam informs me 

Bpicuousl; In general plumage, and 

that with young birds I th p e- 

MOU9 year of both spe m 

p olam, which is pure whit* in 

lared on Apr I 'ord, 1869 th 

th male end gieyish-while in the 

uas no difference in th d 1 p 

f mal . Now the young males nt 

meut of the spurs Th spur 

il t ntirely resemble the females, 

howeier weie ai, yet p at d 

nd h re a greyish- white Bpeculnm, 

meie y by sliithl knob 1 

wh (Ji becomes pare white at aa 

tions I presume that I h Id 

41 1 age than that at which the 

have been informed il anydtfi n 

dult male acquires his other and 

in the rate of devekfm n h d 

n. strougly-marked semal A^ 

been observed snbsequen ly 

f DCs: sec Audubon, 'Oraitb* 

" In some other spe i h 

i i 1 BioKK,phy,' vol. ill lei^S, 

Duck family the specul m djff rs 

pp 49 250. 

^D a greater deKree i th wo 


Cbap. VIII. Sexual Selection. 237 

to one sex. wHlst others are common to both sexes. A diffeicnce 
of this kind in the period of development is not so improbable as 
it may at first appear ; for with the Urthoptera, which assume 
their adult state, not by a single metamorphosis, hat by a sac- 
cossion of moults, the young males of some species at flrst 
resemble the females, and acquire theii distinetiTe masculine 
characters only at a later moult. Strictly analogous cases occur 
a,t the suc&issive moults of certain male erostaceans. 

We tiave as yet considered the transference of characters, re- 
latively to their period of development, only in species in a 
natural state, we will now turn to domesticated animals, and 
first touch on monstrosities and diseases. The presence of super- 
numerary digits, and the absence of certain phalanges, must he 
determined at an early embrj oaio period — the tendency to profuse 
bleedii^ is at least coi^nital, as is probably colour-blindness — 
yet these peculiarities, and other similar ones, are often hmited 
in their transmission to one sex; so that the rule that 
characters, developed at an early period, t«Ed to be trans- 
mitted to both sexes, here wholly fails. But this rule as 
before remarked, does not appear to bo neai'ly so general as the 
conyerse one, namely, that characters which appear lat« in life 
in one ses aie transmitted exclusively to the same ses. From 
the fact of the above abnormal pecnliarities becoming attached 
to one ses, long before the sesual functions are active, we may 
infer that there must be some difference between the sexes at an 
extremely early age. With respect to sesually-limitod diseBses, 
we know too little of the period at which they originate, to draw 
any safe conclusion. Gout, however, seems to fall under our 
rule, for it is generally caused by intemperance during manhood, 
and is transmitted from the father to his sons in a much mora 
' marked manner than to his daughters. 

In the yarious domestic breeds of sheep, goats, and cattle, the 
males differ from their respective females in the shapeor develop- 
ment of their horns, fbrehead, mane, dewlap, tail, and hump on 
the shoulders ; and these peculiarities, in accordance with our 
rule, are not fully developed until a rather late period of life. 
The sexes of d(^8 do not differ, except that in certain breeds, 
especially in the Scotch deer-hound, the male is mUch larger 
and heavier than the female; and, as we shall see in a future 
chapter, the male goes on increasit^ in sizo to an unusually Sate 
period of life, which, according to rule, will account tor his in- 
creased sizo being transmitted to his male offspring alone. On 
the other hand, the tortoise-shell colour, which is confined to 
female cats, is ipiite distinct at birth, and this case violates the 
rule. There is a breed of pigeons in which the males alone ars 


23S I'he Descent of Man. Part 1! 

stieaked with bla«k, and the streiiks asa be detected eTon in th: 
nestlings ; but thej become more conEpicuous at each suecesBiVG 
moalt, so that this ease paxtiy opposes and partly supports tlio 
rul& With the English Carrier and Pouter iiigeons, the full 
deTolopraent of tiie wattle and the crop occurs rather late in life, 
and conformably with the rule, these characterB are transmitted 
in full perfection to the males alone. The following cases perhaps 
come within the class previously alluded to, in which both sexes 
have va^ed in the same manner at a rather late period of liic, 
and liave consequently transferred their new characters to both 
sexes at a corresponding late period ; and if bo, these cases are 
not opposed to our rule : — there esiat sub-breeds of the pigeon, 
described by Nenmeister," in which both sexes change their 
colour diuing two or three moults (as is likewise the case with 
the Ahnond Tumbler), nevertheless, these changes, though 
occurring rather late in life, are common to both sexes. One 
Tariety of the Canary-bird, namely the London Prize, ofiers a 
nearly analogous case. 

With the breeds of the Fowl the inheritance of various charac- 
ters by one or both sexes, seems generally determined by the 
period at which such chariictera are developed. Thus in all the 
many breeds in whieh the adult male differs greatly in colour 
from the female, as well -as from the wOd parent-species, he 
differs also front the young male, so that the newly-acquired 
characters must have api)eared at a rather late period of life. 
Oa the other hand, in most of the breeds in which the two sexes 
resemble each other, the young are coloured in nearly the same 
manner as their parents, and this rendtrs it probable that their 
colours first appeared early in life. We have instances of this 
feet iu all black and white breeds, in which the your^ and old 
of both sexes are alike ; not can it be maintained that there is 
something peculiar in a black or whibi plumage, which loads to 
its transference to both sexes ; for the males alone of many 
natural species are either black or white, the females being 
differently coloured. With the so-called Cuckoo But>-breeds of 
the fowl, in which the feathers are transversely pencilled with 
dark stripes, both sexes and the chickens are coloured in nearly 
the same manuer. The laced plumage of the Sebright bantam 
is the same in both sexes, and in the young chickens the wing- 
feathera are distinctly, though impertectJy laced. Spangled 
Hamburgs, however, offer a partial exception ; for the two sexes, 
though cot quite alike, resemble each other more closely than 

'• 'DasGdnza dcr TaubeDzuctt,' pais, ' Le pigeon Tovageur Iklge,' 
18.J7, a. 21, 34. For (Jie case of 186S, p. 67, 
Ifau streaked pt£eDiu, ie« Dr Cha- 


Okap. vm. Sextial Selection. 239 

do the sexes of the aboriginal parentrspecies ; yet they acquire 
their characteristic plumage late in life, for the chickens are 
distinctly jjencilled. With respect to other characters besides 
colour, in the wild-parent epecies and in most of tho domestic 
hreeds, the males alone possess a well-developed comb; but in 
tho young of the Spanish fowl it is laigely developed at a very 
early age, and, in accordance with this early development in the 
male, it is of unusnaJ size in the adult female. In the CJome 
breeds pugnacity is dereloped at a wonderfully early age, of 
which curious proofs could be given ; and this character is trans- 
mitted to both seses, so that the hens, from their extreme 
pugnacity, are now generally exhibited in separate pens. With 
tho Polisii breeds the bony protuberance of the skull which 
supports tho crest is partially developed even before tho chickens 
are hatched, and the crest itself soon begins to grow, though at 
first feebly ;" and in this breed the adults of both sexes are 
charact«rised by a great bony protuberance and an immense crest. 

Pinally, from what we have now seen of the relation which 
esislfl in many natural species and domesticated races, between 
the period of the development of their characters and the 
manuer of their transmission — for example, the striking fact of 
the early growth of the horns in the reindeer, in which both 
sexes hear horns, in comparison with their much later growth 
in the other species in which the male alone bears horns — we 
may conclude that one, though not the sole cause of characters 
being esclusively inherited by one sex, is their development at 
a late age. And secondlj, that one, though apparently a less 
efBcient causo of characters being inherited by both seses, is 
their development at an early age, whilst the seses differ 
but little in constitution. It appears, however, that some 
difference must exist between the sesea even during a very 
early embryonic period, for characters developed at this age not 
rarely become attached to one sex. 

Sammary and eonclading remarks. — From the foregoing dis- 
cn.^ion on the various laws of inheritance, we learn that the 
characters of the parents often, or even generally, tend to become 
developed in the offspring of the same sos, at the same age, and 
periodically at the same season of the' year, in wtiich they first 

" For full particulars and r«- 250, 256. In regard to the higher 

fereacesmall these points respect- animals, tie seiunl differences wbioh 

ing the sereral bteeds of the lowl, have arisen mder domestication ai> 

see' Variation nf Animals and Plants deacrihed in the Hime wort UEdtr 

aadar Domeiticntioc. vol. i, |ip. the hf ad of each speciei. 


240 Ths Descent of Mtu. p,u;t U 

apiKiared in Iho parents. But thesD rules, owing to unknown 
cauMS, are tar from being fixed. Hence during the modification 
of a species, the successive changes may readily to transmitted 
in different ways ; some to one sex, and some to both ; some to 
the offspring at one age, and some to the ofispring at all ages. 
Not only are the laws of inheritance cstrereely comples, bat so 
are the causes which induce and govern variability. The 
TariatioEs thus induced are preserved and accumulated by 
eesual selection, wUich is in itself an extremely cojnples affair, 
depending, as it does, on the ardour in love, the courage, and 
the rivalry of the males, as well as on the powers of perception, 
the laate, and will of the female. Sesual selection will also 
he largely dominated by natural selection tending towards 
the general welfare of the species. Hence the manner in which 
tho individuals of either or both sesea have been affected 
through sesual selection cannot fail to be complei in the highest 

When variations occur late in life in one sex, and are trans- 
mitl*!d to the same 'les at the same age, tho other ses and the 
young are left unmcfdifled Wlicn they occur late in life, but 
are transmitted to both seses at the Kame age, the young alona 
are left unmodified. Variation'!, however, may occur at any 
period of life in one sex or in both, and be transmitted to both 
eoses at all ages, and then all the individualB of the species 
are similariy modified. In the followmg chapters it will be seen 
that all these cases frequentlj occm m nature. 

Sexual selection can never act on any animal before the ago 
for reproduction arrives. From the great eagerness of the male 
it has generally acted on this ses and not on the females. The 
maloB have thus become provided with weapons for ^hting 
with their rivals, with organs for diECOvering and secureJy 
holding the female, and for esciting or charming her. When 
the sexes differ in these respects, it is ajso, as we have seen, an 
extremely general law that the adult male differs more or less 
from the young male ; and we may conclude from this fact that 
the successive variations, by which the adult male became modi- 
fied, did not generally occur much before the age for reproduction. 
AVhenever some or many of the variations occurred early in 
life, tho young males would partake more or less of the charac- 
ters of tiie adult males ; and differences of this kind between 
tho old and young males may be observed in many species of 

It is probable that young male animals have often tended to 
vary in a manner which would not only have been of no use to 
them at an early age, but would have been actually injurious!— 


DiAv. Villi Scxudi Selection. 241 

49 by acquiring bright colours, which would rcndur them cou- 
spicEOua to their encanies, or by acquiring structures, such as 
gr^at horns, wMch would espend much yital force in thei; 
lievelopment. Variations of this kind occurring in the young 
malus would almost certainly be eliminated through natural 
selection. Witli the adult and experienced males, on the other 
hand, the adyantages deriyed from the acquisition of saeh 
characters, would more than counterbalance some exposure to 
danger, and some loss of vital force. >, 

As tariations which give to the male a better chance of \ 
conquering other males, or of Snding, securing, or charming the 1 
opposite ses, would, if they happened to arise in the female, be I 
of no service to her, they would not be preserved in her through 1 
sesual selection. We have also good evidence with domesticated 
animals, that variations of all kinds are, if not carefully selected, 
soon lost through iutercroBsing and accidental deaths. Conse- 
qnentJ J in a state of nature, if variations of the above kind chanced 
to arise in the female Une, and to be transmitted exclusively in 
this line, they would he estremely liable to be lost. If, however, 
the females varied and transmitted their newly acquired 
characters to their oflspring of both sexes, the charaeters which 
were advantageous to the males would be preserved by tiera 
through sexual selection, and the two sexeswould in consequence 
be modified in the same manner, although such characters were of 
no use to the females; but Ishall hereafter have to recur to these 
more intricate contingencies. Lastly, the females may acquire, and 
apparently have often acquired by transfeienoe, characters from 
tho male sex. 

Aa variations occurring late in life, and transmitted to one 
ses alone, have incessantly been taken advantage of and accumu- 
lated through sexual selection in relation to the reproduction of 
the species; therefore it appears, at first sight, an unaccountable 
iact that similar variations have not frequently been accumu- 
lated through natural selection, in relation to the ordinary 
habits of life. If this had ocsurced, the two seses would often 
have been differently modified, for the sake, for instance, of 
Cfflpturii^ prey or of escaping flrom dai^er. Differences of this 
kind between the two sexes do occasionally occur, especially in 
the lower classes. But this implies that the two sexes follow 
different habits in their struggles for existence, which is a rare 
circumstance with the higher animals. The case, however, is 
widely different with the reproductive functions, in which respect 
the sexes necessarily differ. For variations in structure which 
are related to these functions, have often proved of value to one 
eex and from having arisen at a late period of life, have been 


242 The Descent of Man. Part iL 

u'ansmitlBd to one sex alone ; and such variations, thus preserved 
aud transmitted, have given rise to secondary Bexual characters. 
In the following chapters, I shaJl treat of the secondary 
sexual characters in animals of all classes, and shall eodeavoui in 
each case to apply the prineiplfis explained in the present 
chapter. The lowest classes will ddtajn us for a very short time, 
but the higher animals, especially birds, must be treated at 
coiLsidetable length. It should be borne in mind that for 
reasons already assigned, I intend to give only a fow illustrative 
instances of the innumerable structures by the aid of wliich the 
male finds the female, or, when found, holds her. On the otlict 
hand, all structures and instincts by the aid of which the icalo 
conquers other males, and by which he allures or eseites the 
female, will be fully discussed, as these are in many ways the 

n the pTf^orHonal numbers of the two sexes in animah 
belonging to vario-as citMXfS. 

As no one, as far as I con discover, has paid attention to the 
relative numbers of the two sexes throughout the animal 
kingdom, I will here give such materials as I have been able lo 
collect, although they am extremely imperfect. They consist in 
only a few instances of actual enumeration, and the numbers are 
not very large. As the proportions are known, with certainty only 
in znankind, I will first give them as a, standard of comparison. 

J/an.— In England durmg ten years (from 1857 to 1866) the 
average number of children horn alive yearly was 707,120, in 
the proportion of lOi'6 males to 100 females. But in 1857 the 
male births thronghout England were as 105'2, and in 1865 as 
104-0 to 100. Looking to sepaxafa districts, in Euckingham- 
shire (where about 5000 children are annually bom) the mtan 
proportion of male to female births, during the whole period of 
the above ten years, was as 102 8 to 100 whil>,t m N Wales 
(where the average annual births are 12 873) it was at aigh 
as 106-2 to 100. Taking a stiil smaller distrjct viz Eut 
laudshii-a (where the annual births avenge only 7i)9), m ISbi 
the male births were as 114 6, and in 1&62 as only 97 lo 
100; but oven in this small district the average of the 7385 
births during the whole ten years was ab 104 5 to 100 that is m 
the same ratio as throughout England." The proportions are 
sometimes sUghtly disturbed by uaknown causes; thus Prof. 

** • Tweoty-nlnth Annual Repoft In this report (p. lii.) :. sptcisl d& 
•f tlie a^iBti-ar-Qeneral for 18G6/ cennial tablt i> girou. 


CbaP. V'ill. Proportion of ttie &exes. 24J 

Faye states " that id some districts of Norway there has been 
" during a deeemiial period a steady deficieBcy of boys, whilst 
'' in othors the opposite condition has existed." la France 
during forty-four years the male to the femaJe hirths have been 
as 106-2 to 100; but during this period it has occurred five 
timoa in one department, and six times in another, that tho 
female births have exceeded the males. In Eussia the average 
proportion is as high as 108'9, and in Philadelphia in the United 
Slates as 110-5 to 100." The average tor Europe, deduced by 
Bicies from about seventy million births, is 106 males to lOO 
females. On the other hand, with white, children bom at the 
Cape of Good Hope, the proportion of males is so low as to fluetnat* 
daring saccesaive years between 90 and 99 males for every 100 
femalen. It is a sii^ular fiict that with Jews the proportion of 
male births is decideiJly larger than with Oliristians: thus in 
Prussia the proportion is as 113, in Breslan as 114, and in 
Livonia as 120 to 100; the Christian births in these conntries 
being the same as usual, for instance, in Livonia as 104 to lOO.™ 
Prof. Faye remarks that " a stiU greater preponderance of 
" maJes would be met with, if death stmok both seses in equal 
" T>roj)ortion in the womb and diurii^ birth. But the fact is, that 
" for every 100 still-born females, we have in several countries 
" from 134-6 to 144-9 still-born males. During the first four or 
" five years of life, aliio, more male children die than females ; 
" for example in England, daring the first year, 126 boys die for 
" every 100 gu:!s— a proportion which in France is still more 
" unfavourable.™' Dr. Stockton-Hough accoaats for these facts 
in part by the more frequent defective development of males 
than of females. We have before seen that the male sex is more 

" For Norwuy and Roaiia, Me 

343. Dr. Stark also remarks 

abstract of Prof. Faje's reseu-cihes, 

(■Tenth Annual Report of Births, 

in 'ISiitiih and Foreign Medioo- 

Deaths. &c., in Scotland,' 1867, p. 

Chirure. ReviBw,' April, 1867, pp. 

ijviii.) that " These eiaraples may 

.543, 345. Kor France, the ' An- 

" aulfi;:e to shew that, at almost 

nuiii™ ponr I'An 1867,' p. 213. 

" every stage of life, the males in 

For Philadelphia, Dr. Stockton- 

" Scotland have a greater liabilitj 

Hough, ' Social Science Assoc.' 1874. 
For the Cape of Good Hope, QueMlet 

" to death and a higher death-rate 

'■ than the females. The fact, how- 

as quoteiJ bj Dr, H. B. Zoutevecn, 

"ever, of this pecnliarity being 

in the Dutch Translation of thin 

" most strongly developed at that 

work (vol. i. p. 417), where much 

" infantile period of life when th« 

infoimation ia given on the propor- 

" dress, food, and general treatmeDt 

tion of the wies. 

" of both eeies are alike, seema tc 

" In regard td the Jews, see M, 

" prove that the higher maledealh- 

rhurv, 'La Loi de Piodnctioa des 

.. ......;,— i'' ^^\..,\t^ 4„. ., 


244 The Dest-ent of Man. Jab* IL 

VEiriable in structnre than the female; and variations iu im- 
portant organs would generally be injurious. But the size of 
the body, and especially of the head, being greater iu male tiaii. 
lemole infiinta is another cause; for tJie males are thus more 
liable to be injured during parturition. Consequently the still- 
born males are more numerous ; and, as a highly competent judge, 
Dr. Criohton Browne," bellevea, male infanta often siiffer in health 
for some years after birth. Owing to this excess is tte death- 
rate of male children, both at birth and for some time sub- 
sequently, and owing to the esposnre of grown men to various 
dangers, and to their tendencj' to emigrate, the females in all 
old-settled countries, where statistical records have been kept," 
aio found to preponderate considerably over the males. 

It seems at first sight a mysterions fact tbat in different 
nations, under different conditions and climates, in Naples, 
Prussia, 'Westphalia, Holland, France, England and the United 
States, the excess of male over female births is less when they 
are illegitimate than when legitimate." This has been explained by 
difierent writers in many different ways, as from the mothers 
being generally young, from the large proportion of, first preg- 
nancies, &c. But wc have seen that male infante, from the largo 
size of their heads, suSei more than female infants daring 
parturition ; and as Iho mothers of illegitimate ehildren must be 
more Uable than other women to undergo bad labours, from 
various causes, such as attempts at concealment by tight lacing, 
hard work, distress of mind, &c., their male infents would 
proportionably suffer. And this probably is the most efBoieut 
of all the causes of the proportion of males to females bum 
alive being less amongst illegitimate children than amongst the 
legitimate. With most animals the greater size of the adult 
male than of the female, is due to the stronger males having 
conquered the weaker in their straggles for the possession of 
the females, and no doubt it is owing to this fact that the two 
seses of at lea^t some animals differ in sizs at birth. Thus 

" ' West Riding Lunatic Asylum P.iraguay, actording lo the in:i:urac 

Reports,' vol. i, 1871, p. 8. Sir J, Azara (' Voyages dans I'Am^-iilu 

SimpsQn has proved that the ^-■^ ■*' • — -' '""^ - "■■' """ 

of Che male infant Bioe«ds ti 

1809, p. 60, 179% 
.oe«as mac ui ms women an to the men in ths 
of an inch in proportion of U to Vi, 
by l-8th in » Babbi^e, ' Edinburgh Jouru;;! 

Quetelct has of Soience,' 1829, vol i, p. 88 ; ai^i-. 
children. On 

than mani see Dr. Danran, 'iV illegitimate children in Engh 

cnndity, Fertility, Sterility,* 1871, lee -Report of Regis ttar-Grm 

f, 382. "■— ^o^-:- 

" Witt the savage Giiaraot3 DJ 


Chap. VII!. Proportion of the Sexes. 245 

we havo the curioiis fact that we may attriliuto the more 
frequent deaths of male than female infants, esijeciaJly amongrt 
the illegitimate, at least in i)art to sexual eeleetion. 

It has often been supposed that the relative age of the two 
]iarents determines the sei of theoflspring; and Prof Leuokart'" 
has advanced what he considers sufficient evidence, with respect 
to man and certain domesticated ammaJs, that this is one impor- 
tant though not the sole fiictor in the result So ^;aia the period 
of impr^uation relatively ta the state of the female has heen 
thought by some to he the efficient cause ; but recent observa- 
tions diaoountenance this belief. According to Dr. Stoctton- 
Hough,** the season of the year, the poverty or wealth of the 
parents, residence in the country or ta citieB, the crossing of 
foreign immigrants, Ac, all influence the proportion of the 
sexee. With mankind, polygamy has also been supposed to lead 
to the birth of a greater proportion of female infants ; but Br. J, 
Campbell " carefully attended to this subject in the harems of 
SJam, and concludes that the proportion of male to female births 
is the same as from monogamous unions. Hardly any animal 
has been rende.isd so highly polygamous as tie English race- 
horse, and we shall immediately see that his male and female 
offipring are almost exactly equal in number. I will now give 
the facts which I have collected with respect to the proportional 
numbers of the seses of various animals ; and will then briefly 
discuss how far selection has come into play in determining tho 

UnreeE.— Mr. Tegetmeier has been so kind aa to tabulate for roe fmm 
the ' Eaoing Calendar' the birtbii of race-horees durin" n period of 
tBenly-ono yeara, viz., from 1846 to 1867; 1849 being omitted, as no 
returns were that yeiir published. The total births were 25,560," coa- 
aisting of 12,763 maJea and 12,797 females, or ia the proportion of 99-7 
melaB lo ] 00 females. Ah (hese Qumhera are tolerably iarge, and sa 
they are druwn from all parts of England, dining several years, we may 
with mnoh confidence conclude tl^t with the domiretio horee, or at 
least with the reoe-horae, the two seses are produced in almost equal 
sumbeiB. The fiuctuations in (he proportions duihig successive yearx 

" Leuckart (in Wagner ' Hand- notice, as shewinj how infertile 

w6i-terbuoh dor Phys.* L, iy. 1853, these highly-nnrtuted and rather 

1. 774. ulosely-intu'bi'ed animata have be- 

" Social Science Asbm. of Fhila- come, that not far from one-third o! 

delphia, 1B74. . the ma.Bs &iled .s produce 

" ' Antliropoli^cal Rfiew,* foals, Thm during 1866, 809 moli; 

April, 1870, p. QTia. ■ collB ^nd 816 female colta were bora, 

*■ During eleven yeai's a recorl and 743 mares failed to produw 

was tept of the number of mares offspring. Duiing 1867, 836 malea 

which prored barren oc prematurely and 902 fbmales were bom, apd 7tH 

illppcd their foals; an^ it desetres wares failed. 


246 "the Descent of Man. Faei H, 

are oloBclj like thofe whicli 0(^ClIr n'ttli mankinrl, ivbeo a dma,ll acJ 
thmlj-populattni area ia contddered ; t)jas in 1S56 the male tiorses were 
aa 107-1, and in 18S7 aa only 92'6 to 100 females, lu the tabulated 
retarns tbo ptvportionB ¥»ry in cycles, for the males exceeded the 
females duriii); six socceaeive years ; and (bo females exceeded the 
nmles during two periods casli of four years: this, however, may lie 
aooidontal : at least J can detect nothing of the kiiid with man in the 
deceniiial table in the Begiatiar's Report for 1866. 

Dogt. — During a period of twelvH years, from 1857 lo 1868, the birtlis 
of a Ih^^ umabBX of greyhounds, tiuxjughoat England, were sent to 
tbe ' Field' newspaper ; and I am a^n iudebtod to Mr. Tegetmeier for 
OBrefully tabulating the results. The recorded biiths were 6878, 
cunsititing of 3605 males and 3273 l^enjiiles, that is, in the propurtiun of 
110-1 maTeB to 100 females. The greatest fluotUHtions oeourrcd in 
1864, when the proportion waa as 95'3 males, and in 1867. as 116-3 
midea to 100 femiiles. The above average proponioD of IlO'l to tOO is 
prubflbly nearly ocutect in the case of iho greyboimd, but wtotlier it 
would hold with otbet domesticated breeds is in some dwree doubtful. 
Mr. Cupples has etiqnired from severul greet breeders of dogs, and finds 
that all without exception believe that females are prodnt'ed in excess ; 
but he suggests that this belief may have arisen from females being 
less valued, and from tbe consequent disappointment prodneing a 
stronger impre«sion on the mind. 

Sfti™. — The sexes of sheep are not aacertainid by agricnlturista ontil 
several months aftor biitb. at the period wbun the mules are castrated i 
so thnt the following returns do not give the proportions at birth. 
MoTBoyec, I find that eeverai great breedum in Scollanii, who annnally 
raise some thousand sheep, are firmly oonvinced that a larger pro[«rtioii 
of males flian of femali:s die duriiiK the first year or two. Therefore the 
proportion of males would bo somewhat larger at birth than at the uge of 
castration. This is a remarkable coindileuce with what, as we lave 
seen, occurs wilh mankinil, and both cases probably depend on tbe 
aume caose. I have received returns from four gentlemen in England 
ivho have breil Lowland sheep, chiefly Leioesters, darling the last ten to 
sixteen years; they amount altogelher to 8^65 births, oonsisting of 
44U7 males and 4558 t'euiales; thai is in tbe proportion of 1)6 7 mal^ to 
100 females. With respect to Cheviot and blacb-faccd sheep bred in 
Scotland, I have received returns from six breeders, two of them on a 
>irge scalp, chiefly fur the years 1867-1869, but some of the returns 
extend back to 1862. The total number recorded amounts to 50.685, 
consibtiug of 25.071 males and 25,611 females, or in the proportion of 
97.9 males to 100 females. If we take the English and Scotch retmns 
together, the total number amounts to 59,650, oonaiating of 29,478 
males and 30.172 females, ur as 97'7 to 100._ So that with sheep at the 
age of castration the females are certainly in excess of flie males, but 
prubably this would not hold good at Ijirth." 

Of (hitU I have reotived returns fiom nine gentlemen of 982 births, 
too few to be trusttd ; these consiated of 477 bull-calves and 505 eow- 

" I am much indebted to Mr. tion to the premature deaths gf the 

Cnwles for having procured for me males, — a statement eubsequently 

the above returns from Sootliind, as confirmed by Mr. Aitohison and 

well HE Eome of the folluirtng re- others. Tl> thii latter gentleman, 

turns on tattle. Mr. B. Elliot, of and to Mr. Payan, 1 owe my thsnlu 

Laighwood, first called my atten- for Urje returns as to sheep. 


Chap. VIIl. Proportion of the Sexes. 247 

calves; i.e., in the pcoportion of 9i*4 males to 100 fejoalus. The Rtv. 
W. D. Fox informs me that in 18G7 out of at ealvea bom on a farm in 
Derbyabire only ntie was a bulL Mr. Harrison Wfir has enquired from 
Beveral breeders of Pig», and moat of them estimate the male to the 
female births ad aboDt 7 to G. This same gentleman hue bred Eaiihiiii 
for many yoara, and haa noticed that a far great<jr number oi' buoka are 
produced than does. But estimations are of little valoe. 

Of maoimalia in a state of nature 1 have been able to learu very 
Jittle. In regard to the common rat, I hase received coafliotinp 
Btatement9. Mr. B. Elliot, of Laighwood, informs me that a ratn^lcher 
assured him that be had always found the males in great o^ceas. BTen 
with the youug in the oesi. In oonseqaence of this, Mr. Elliot 
himself subsequently exLunined some hundred old ones, and foand Iho 
etutement true. Mr, F. Bookland has bred a large number of white 
rata, end be also belieTea that the males greatly ex{^eed the females. 
In regard to Moles, it is said that " the males are much more numeroua 
" than the females i"** and as the catching of these animals is a special 
occupation, the statement may pethapa be trusted. Sir A. Smith, in 
desoribiug an antelope of S. Afiiea" iKvbne dliprtprymnui), remarks, 
that in the herds of this and other species, the males ate few in number 
compared fvitli the ftauales : the oativea believe that they are born in 
this proportion ; others believe that the younger males are cxpelleii 
from the herds, and Sir A. Smitb says, that though he haa himself 
uever seen herds oonsiating of young males alone, otbers afarm that 
this doea occur. It appears probable that the young Vihon expelled 
from the herd, would -rfleu fall a prey to the many beasts of preyof tho 


With respect to the Fowl, I have received only one account, namely, 
that out of 1001 cbiokens of a highly-bred stock of Cochins, reared 
duricg eight years by Mr. Streteh, IS7 proved males and Sll females: 
i.e.. as 9t-7 to 100. In regard io domeafic pigeons there is good 
Bvidanca either that the males are produced in exeesa, or that they live 
longer ; for these birds invariably pair, and single males, as Mr. Teget- 
lueier informs me, can always be purchased cheaper than females. 
Usually the two birds reared from the two eggs laid in the same nest 
are a male and a female ; but Mr. Harrison Weir, who has been so large 
H breeder, says that he has often bred two cocks ftom the same nest, 
and seldom two heos ; moreover, the hen is generally the weaker of the 
two, and more liable to petish. 

With respect to binla in a state of nature, Mr. Gould and others" 
are couvmced that the males are generally the more nnmerous; and 
aa the yonng males of many species resemble the females, the latter 
would naturally appear to be the more nmneroua. Large numbers of 
pheasants are reared by Mr. Bakerof Leadenhall from egga laid by wild 
birds, and he informs Mr. Jernier Weir that four or five males to one 
female are generally produced. An experienced observer rpmnrks," 

*• Bell, 'History of British Quad- iv. s. 990) comes to the same eon- 

ropeds,' p. 100. elusion. 

" ' Illaetrationa of the Zoology " On the aothority nf L. Lloyd, 

of S. Atrioa," 1849, pi. 29. 'Game Birdi of Sweden," 1867 fp 

*' Brehm (' IUds',. Thieilebeu,' B. 12, 132. 


248 The Descent of Man. Piicr II, 

lliat in SeaodinaTia the broods of the oaperaajlrie and blaok-ccck 
coDtuin more males than, females ; e,Qd that nith the Dal-ripa (a Mod 
of ptHTmigHu) mora males than females attead ths Ma or placea ut 
oanrtehip; but this latter ciroumeliuioe is accounted foi by some 
chaeryers by a greater number of hen birds being iiilled by vermin. 
Prom vacious &cis given by Wliite of Selbome," it seems clear that 
the males of the partridge must be in considtrable escess in the Boulh 
of Kngland ; ani I have been assured that tbis is the case in Scotland. 
Mr. Weir on enquiring from the dealers, who rfoeiTe at cert^n seasona 
largo nmnbeis of ruQs {Ma/ihstet pugnox), 'was told that the males aro 
much the more numerous. This same naturalist has also enquirod for 
me from the birdcatchera, who annnally catch an astoaisiiing nmabft 
of varinus smaU species alive for the London tnarktt, and he was un- 
hesitatingly nnsneml by an old and truscworthy rano, that with Ibo 
lihoffiiich tbe males are in large esoess; he thought as high as 2 males to 
1 female, or at least as high as 5 to 8." The males of tlie blackbird, 
he likewise maintained, wore by iar the more numerous, ivhetVier 
caught by traps or by netting at night. These statements maj' 
apparentjy be trusted, because this same maTi said that the Si 
about equal with the iark, the twite {lAwttia, ituintitflct), and g 
On the other hand, he is certain that with the common linnet, tnu 
femalea preponderate greatly, but anequaJly during diiferent years ; 
during some years lie has found thefemak-a to the males as fonr to one. 
It bhouid, however, be borne in mind, that the chief seiison tbr catcliing 
birds does not begin till September, so that with some spoeies partial 
migmtioas may have begun, and the flncks at this periid often oonsist 
of hens alcne. Mr. Sulvm paid particular attention to the sexes of the 
bummiDg-birds in Central America, and he is convinced that with 
most of the species the males are in excess; thus one year he poooted 
204 specimens belonging lo ten species, and these consisted of 166 
males and of only 3S females. With two other species the females were 
in eicesa ; but the proportions apparently vary either during different 
seasons or in different localities; for on one occasion the males of 
Canroijiopferiu ftemfleufunis were to Uie females as 5 to 2, and on 
another occasion" in exactly the reversed ratio. As bearing on this 
latter point, I may add, that Mr. Powys found in Corfu and Bplms 
the sexes ot the chaffinch keeping apart, and " the females by for the 
" most numerous ;" whilst in Palestine Mr. Tristram found " the male 
" flocks appearing greatly to esoeed the female in number."*' So 
again with the Cuiaeai'is maj'irr, Mr. G. Taylor" says, that in florida 
there were " very I'ew females in proportion to the males," whilst in 
Eonduraa the proportion was the other way, the species there bavin)? 
the charaitcr of a polygamist 

" 'Nat. Hist, of Salbornc,' letter Brer caught by one in a single 

nil. edit, of 1825, vol. i. p. 139. day was 70. 

" Mr. Jenner Weir received •■ ' Ibis,' vol. ii. p. 280, as qutttd 

similar intormation, on making en- ra Gonld's ' TroohiUdfe,' 1861, p. 

quiries during the following year, 52. For the foregoing proportionfi, 

To shew the nnmbtr of living chaf- I am indebted to Mr. Salvin for a 

finches caught, I may mention that table of his results, 
in 18S9 there was a match between " 'Ibis,' 1860, p. 137; and 1887 

two eiperts, and one man caught p. 369. 
in a day 62, and another 40, mi^le " 'IWs,' 1862, p. 1^7, 


Proportion of tlie Sexes. 


WiUi Fish the proport ou'J numbera of the Eoxes ran Tie OBPeifBiiiod 
only by oalching IhelO in the aJ It or nearly ailult state ; and there 
ore many diffloalties m arriving at any juat conoluaion.'* Infertile 
fomalea might readily be mistuken for males, as Dr. Giinther bus 
remftrked to me in reeard to trout ft tb eorae spaciefl ibe malea are 
beliered to die soon after fertilising the ota. Witli many speoioa tl-e 
malBB are of much Bmailei size than the females, so that a large 
camber of malea would escape from the same net by which tlie females 
were caught M. Carbnnnier," who has especially attended to [ho 
naturalhiatury of thepike (Eioxlwiint), statea that many males, owing 
to their small size, are devoured by the larger females ; and he believer 
that Ibo males of almost all fish are exposed from Ibis seme canae to 
greater danger thau the females. Nevertheless, in tlie ffw f»BeB in 
whioh the proportional numbera have been actually observed, the 
malea appear to be largely in exceaa. Thus Mr. It. Baist, the superin- 
tendent of the Stormontfleld esperiments, aays that in 1865, out of 70 
salmon first landed for the jiurpose of obtaining Iha ova, upwards of 60 
were malea. In 1867 he again " oalla altentioii to the vast dispropottioo 
" of the malea to the females. We had at the outset at least ten males 
"to one female." Afterwards females sulHcieut for obtaining ova were 
procured. He adds, " from the great proportion of thn males. Ihey aro 
"constantly Sghting and tearing each other on the spawning-beds."" 
Thia disproportion, no doubt, «an be accounted for in part, but whether 
wholly ifl doubtful, by the males aacending the rivers before the 
females. Mr. F. Buokland remarks in regard tn trout, that " it is a 
"' curious fact that the males preponderate very largely iu number over 
" the femalea. It iniiariahlv happana that when the firit rush of fish is 
" made to the net, theie will he at least seven or eight males to one 
"female fonnd captive. 1 cannot quite account for this; either tht 
" males are more numerons than the females, or the latter seek safety 
" by concealment rather than flight." He than adds, that by carefully 
8e!i.rohing the banks sufflciant femalea for obtaiiiing ova can be found.'' 
Ml-. H, Lee infomis me that out of 2L2 trout, taken for this purpose in 
Lord Portemouth's park, 150 were malea and 62 leinalos. 

The males of the CTOrinidiB likewise aeem to be in excess; but 
aeveral members of tins Family, viz., the carp, tench, bream' and 
minnow, appeal tBgnlarly to follow the practice, rare in the animal 
kingdom, of polyandry ; for the female wMUt spawning is Iwa a 
attended by two males, one on each side, and in the Oas f the b a n 
by three or four males, Thia fact is bo well known, tl at t s alna s 
reeommended to stock a pond with two male tenches t one f male, 
at least with three malea to two females. With he mirni w nn 
excellent observer states, that on the spawning-beds tl malea a e ten 
tiiBos as numerous as the females; when a female comes am ngst the 

" Lenciiart quotes Bloch (Wag- 18. 1869, p. 369. 

Uer, ' HandwiirtCTboch der Phj^s.' " 'The Slormontfield Pisdcnl- 

B. iv. 1853, a. 77a), that with fish (nral Eiperiments,' 1866, p. ^3. 

(l-o™ oTo tiu!/.= •,. many males as The ' Field' newspaper, June 29tii, 


The Descent of J 

males,"' she is immeijiately preesed closely by a male on eftcli Bide- 
"and whon they have bsen in tbat situation foe a time, ars «upeceeded 
" by olljer two males." " 


In this great Clasa, the Ijepidoptenv almost alone afford means for 
Juiiging of the proportional numbera of the BWtea ; for Oiey have beeu 
tolleeted with spefiial care hy many good observers, and have been 
lai^ely bred from the egg or catetplllar state. I had hoptil that isome 
breeders of silk-moths might have kept an esaet record, but after 
writing to France and Italy, and ooQ.sulting variouB treatises, I cannot 
lind tbat this has ec^ beeo done. The ^neral opinion appears to be 
tbat tlte sexes ore nearly equal, but in Italy, as I bear from Profesaor 
Caneatrini, many breeders ere convinced tbat the females are produced 
in tscesa. This same oaturalist. however, infurms me, that in the two 
jfearly broods of the Ailauthus silk-moth (Ban^yx, eyalMii), the males 
greatly preponderate in the first, whilst in the second the two sexes are 
nearly equal, or the females rather in excess. 

In re!;ftrd to Butterflies in a state of nature, several observers haTS 
been much struck by the apparently enormous pioponderance of the 
males.''' Thus Mr. Bates," in speaking of several species, about a 
htindrod in number, which inhabit the Upper Amamna, says tbat tha 

maies are iiiucli more numerous than the females, even in the propor- 
tion of a hundred to one. In North America, Edwards, who had great 
experience, estimates in the genus Papilio the males to the females oa 
tour to one ; and Mr. Walsh, who informed me of this statement, says 
that with P. turniM this is certainlj^ the case. In South Africa, Mr. B. 
Trimen found the males in excess in 19 species :" and in one of these, 
which swarms in opin pluoes, be estimated the nnmber of males ea 
fifty to one female. With another species, in which the males are 
numerous in certain localities, lie collected only five females during 
aoTfn years. In the island of Bourbon, M. Maillard states that the 
males of one species of Fopilio arc twenty times aa numeruus as the 
females." Mr. Trimen informs me that as fet as he has himself seen, 
or heard from others, it is rare for the females of any butterfly to 
exceed the mules in number; but three South African species per- 
haps offer an exception. Mr. Wallace" states that the females of 
iyridthogttra, eriBsui, in the Malay archipelago, are more common and 
more easily caught than the malw ; but this is a rare butterfly. I may 

" Yariell, ' Hist. British Fiabes,' 
(.il.i.l8i6,p.307; rathe Cyyrinns 
carpio, p. 331 ; on the ISnaa valgar '- 

p. liSl ; on the Abntmia brama, p. 

aona,' vol. ii. 1863, p. 228, S47. 

33G. Bee, fiir the minnow (Z™- 

" Fonr of these cases are given 

fliKUJ plioxmus), ' Lopdon's Mag. of 

by Mr. Trimen in bis ' Khopalocera 

Kat. Hkt.' vol. T. 1832, p. 682. 

Africa Auativilis.' 

" Leuckart quotes Meineofce 

" Qnoted by Tiimen, 'Transact. 

(Wagner, ' Handworterbncli dec 

Ent. Sot' vol. V. part iv. I8G6, p. 330, 

I'hys.' B. it. 1853, a. 775) that 

" 'TraHBHBt. Lian. Soc.' vol. ixv. 

tb* male? jf Battsrfliej .ire three 

p. 37. 


Proportion of the Sexes. 

bare add, that io Hypcryflita, a genua of moiha, Gncn^ says, that 
fi-om fonr to £fs females are sect iu collections from ludlu for ooa 

When this flubject of ttie proportional numbera of the seies of inseetS 
waa brought before the Entomologioal Society," it was generally 
tulmitted that the moles of most Leprdoptetci, hi the adult or imago 
state, are cauelit in greater numbers than the feraalea: but this &at 
was attributed by Tarioua obser?era to the more retiring habits of tha 
females, and to the males emerging earlier from the cocoon. This 
latter ciicumatance is well known to occur with most Lepidopters, as 
well as with other iasects. So that, as M. PerEonnat rcmajrka, the 
males of the domesticated Soinbips Yammaai, are Useless at the b^t)- 
Ding of the aeaaon. and the females at the eud, fiom the want of 
mates.'" I cannot, however, persuade myself that tiiese oansea suffice to 
explain the great escesa of males, in the above caEfis of certain butter- 
flif« which are estremelj common in their native countries, Mr. 
StaintoD. who has paid very close attention during many years to the 
Bmaller moths, informs me that when he collected them in the imago 
state, ho thought that the males were ten times as numerous as the 
females, but that since he has reared them on a kii^ scale from tbe 
caterpillsi state, he is convinced that the females are the moie 
numerous. Several entomologists concur in this view. Mr. Double- 
day, however, and some others, take an opposite view, and aie eon- 
viuced that they havS reared from the egga and eaterpiUara a larger 
proportion of males than of females. 

Besides the more active habits of the males, their earlier emerging 
&oin the cocoon, and in some cases their frequenting more open 
stations, other canses may be assigned for an apparent or real diflia'ence 
ib the proporfional numbers of the aeiea of Lepidopl«ra, when cap- 
tured in the imago state, and when reared from the egg or caterpillar 
state. I hear from Professor Canestriui, that it is believed by many 
btetdets in Italy, that the female CBterpillar of the silk-moth suffers 
more from tbe recent disease than the mate ; and Dr. Standinger 
informs me that in rearing Lepidoptera mure females die in the 
cocoon than males. With niany species the female caterpillor is lai^r 
then the male, and a collector would naturally choose tbe Unest 
specimens, and thus unintentionally collect a larger number of females. 
Three coUectors have told me that this was their practice; but Dr. 
Wallace is sore that most collectors take all the apeoimens which tbey 
can find of tbe rarer kinds, which alone are worth the trouble of 
reariug. Birds when surroanded by eaterpiUara wonld probably 
devour the largest ; and Professor Canestriui informs me that in Italy 
some breeders believe, though on iasuffident evidence, that in the %iA 
broods of the Ailanthua silk-moth, the waapadoatroj a larger number of 
the Eemale than of the male calerpiilars. Dr. Wallace further remarka 
Uiat female caterpillars, from being larger than the males, require 
more time for their development, and consume mme food and mots 
ture; and thus they would he espoaed during a longer time to 
danger from iohneumona, biidti, &c., and in times of scarcity would 
perish in greater numbers. Hence it appears quite possible ihsil 

'• 'Pmc Entomolog. Soc' Feb. 'Proc Eat. &oc." 3ni poiies, »o). v 
I7th, 1868. msT, 1.487. 

'• Quol*d by Dr. Wall^e ib 


The Descent of Man. 

t f t fw fml 

Tj pid pt m J -a li maturitv 

th m 1 on I f pe 1 

bjwt w eem d w tb (heir 

rel t uml t jn t tj wh 

th aJyt popagate 

111 ni wh h Ih m-U 

f •i:\;&ai m th ™ "T gate in 

1 d ry Dih >d 

gl f mal ppi tly d :aW8 a 

gr t ess fm 1 th gh th 

t t m y p h ps b acoo nted for 
m lea from fli u coc «) . Mr. 

by h aril m f th 

fctaintn C nna lo th t from tw 

1 totwtyml my Ren be 

fe ooggtdrod fml 

HaAwforu/Mi It is wt^U 

imwnthtl s t Of } q rv Sim carpi-i 

b posed caj, at mhers f m lea lleet ro d h end if 
tonfln&l in a, ronm will e\f,ti tome down the ihimney to htr Mr 
Doubleday b I evea that he boa seen from fifty lo a hundred males of 
both lho~e "peiieH atiracted in the envir'e of B flingla day bt a femnle 
in confinomect In tbe Isle of Wight Mr Inmen Cipoaed tt t" i 
in whi(,h a female f the LaaocEimpa Lad b fn oonhncd on the 
previoHs day and fiie males booh endeavoured !o gain admittaiic« 
In Austria M \erruau5t, hanng placed the female ot a email 
Bombyx m a bos ra hia (jotket, was followed hi a crowd of male^ so 
that abiut 2iH) intored the house with him " 

Mr Doubleday baa called my attention to M. Siaudingera" hst 
of Lepidoptera, which gives the prices of the males and femalee of 
30O speciua or well-morEed vaiietifa of butterflits-CKhopalofeia). The 

but in IH of the rater speoiea Ihey differ; the males being in all csaes, 
eicepting one, the cheaper. On an aversge of the prices of the 113 speciea, 
the price of the male to that of (he female is as 100 to 149 ; and Ihis 
apparfntly indicates that myeraelv the males exceed the females in 
the aanie proportion. About 2000 apeoiea or rarielJeB of motha 
(Hetetoceca) are catalogued, those with wingless females being here 
excluded on account of the difference in habits between the two sexes ; 
of these 2000 apeciea, 141 difffer in price according to pes. the malea 
of 130 being cheaper, and those of only 11 being deaier than ihe 
females. The average price ot the males of the 130 species, to that of 
the females, is as 100 to 143. With respect to (be butterflies In this 
priced list, Mr. Donbleday thinks (and no man in England has had 
more experienceX *'!"•' tbere is nothing in the habits ot the species 
which can account for the difference in the prices of the two sexes, 
and that it can be accounted for only by an excess in the number of 
the males. But I am bound io add that Dr, etaudii^er informs ine, 
that he is himself of a different opinion. H" thinks that the less active 
habits of the fcnmles and the earlier emergence of the males will 
BCtount tor hia collectors seooring a larger number of males than 
ottemales, and oonsequently tor the lower prirea of the former. With 
respect to specimena reared from the caterpillar-state, Dr. StandiDgBr 
believes, aa previously stated, that a greater nnmber of females than ol 
males die whilst confined in the cocoons. He adds that with certain 
species one sex seems to prepondemte orer the other during oertaiB 

Of direct obscrvntions on the aexcs ot Lepidoptera, reared either 

" Blacohard, ' Metamorphoses, " ' t.epidopteren - DonblslteB 


Prt'porttoH of the Sex^i. 


from e^^ or catei|iillara, I hare received only the few followiog 

The Rev. J. Hellins" of Eieter reared, daring IBSS.ll 

101^03 of73 species, which coDsistod of. . ./[ 
Mr. Aibert Jones o[" Elthain reared, during 18G8,l| 

imagos of 9 species, which consistsd of . . .y. 
DnrinS 1869 ha reared Images from 4 spedes, con')| 

siBting of i 

Mi. Buckler of Emsworth, Hants, dnring 1869,1 

reared imagos from 74 species, consistiflg of. , / 
Dr. Wallace of Colchester reared from one brood ofl 

Bomhyi crnthia / 

Dr. Wallace raised, fratu cocoons of Bombyi Pernyil 

sent from China, during 13S9 / 

Dr. Wallace raised, dnring 1K68 and 1869, freia twu\ 

lots of oocoom of Bombji yama-mai . . ,/ 


So tbat in these eight lota of cucoohb and eggs, males were prodaoad 
in OKoeBS. Taken together the propottinn of males is as Xt'l'l 
to 100 femeles. Bat the nombera are hardly large enongh to be 

On the whole, from these various sources of evidence, all pointing 
in the same direction, I infer that with moat Epocies of Lepidopteis 
the mature uuilea generally exceed the females ic number, whatever 
the proportiona may be at theii first emergence from the e^. 

With reference to the other Orders of insects, 1 have oeen aWa 
to collect very little reliable informatioD. With the atag-beetle 
(taeonin cernis) " tlie malea appear to fae much more noioeroua 
"than the femalea;" hut when, as Cornelius remarked during 1867, 
an unusual number of these beetles appeared in one part of Germany, 
the lecaalea appeared to exceed the u^es as six to one. With one of 
the ElutoridiB, the males are said to be much more numerona than the 
females, and " two or three ore often fonud united with one female ;" 
'■ HO that here polyandry seems to prevail." With Siagonium (Staphy- 
liuidie), in which the males are furnished with horns, " tlie females are 
*< far more nnmeicns than the opposite sex." Mi. Janson stated at the 
Entumological Society that the females of the bark- feeding Knniciu) 
viUoeui ace so common as to be a plague, whilst the males ore so rarn 
u* to be hardly known. 

" This naturalist 

has been ao "* Giipther's ' Record ol 

soma resniu logical Liteiature,' 1^67, j 

n which the Ob the eiceas of female L 

preponderate; ibid^p. 250, On the males o 

h"-ures were EUa in England, Westwood, ' ! 

3 t ni ibic Class, of InseotB,' vol. i, p. 1! 
the SiagoTiiura, ibid. p. 173, 


254 tlie Desceid of Man. Par! T!. 

It iH Lardly worth whils spying tuijthing aliout the proportion it 
the eeses in certain species and even groups of insects, for Ibe ujnl.i, 
are miktiann or rei; rnre, ami the femalea are parthenogem^c. tli» 
fa, ieitile vithout sexual unioo; examples of this are afi'orded bj 
•Bveral of the Cynipiiiee.'^ In al! the gall-makiog Cynipidie known 
to Mi. Walsh, the females are four or live times us numerous ax the 
males ; and so it is, as he informs me, with the gall-mating (Jeoidomyiiie 
(Diplera). With sorae common species of Saw-flies (Tenthrediiue) 
Mr. F. Smith has reared hundreda of specimens from iarvie of all 
sizes, but hac never reared a single male : on the other hand, Curtis 
says,'' that with certain species (Athalia), bred by him, the males were 
to the fomoiefi as six to one ; whilst e;sactly tbe reverse occurred with 
the mature insects of the ssme species caught in the fields. In the 
family of Bees, Hermann Muller," collected a large number of 
specimens of many ajiecics, and reared others from tl:e cocooas, and 
counted tbe seses. He found that the males of some species greatly 
exceeded the females in number; in others the reverse occurred; and 
in others the two sexes were nearly equal. But as in most cases the 
mules emerge from the cocoons b^ore the females, they are at tbe 
commencement of the breeding season practically in eicesa, Miiiier 
also observed that the relative number of the two sexes in sume 
species differed mnch in different localiUes, But as H. Miiller \\&» 
himself remarked to me, these remarks must be received with 
some caution, as one sex might more en^Uy escape observation than 
the other. Thus bis brother Fritz Miiller bas uatii.>ed in Brazil that 
the two sexes of the same species of bee sometimes frequent different 
kinds of flowers. With respect to the Orthopteca, I know hardly 
anything about the relative number of the sexes : KLorte," however, 
says that out of 500 locusts which be examined, the males were to 
the females as five to six. With the Nearoptera, Mr. Walsh slates 
that in many, but hy no means in all the species of the Odonatous 
group, ihei-e is a great overplna of males ; in the genns Heta^rina, also, 
the males are generally at least four times as nmnennia as the femslcs. 
In certain species in tlie genus Glomphus tbe males are equally in 
excess, whilst in two other species, the females are twice or tbrice 
OS numerous as the malea. In some European species of Fsocua 
thousands of females may be collected without a siugle male, whilst 
■mth other species of the same genus both sexes are common." In 
England. Mr. MacLacblan has captured hundreds of the female 
A'palania lavMebrit, but lias never seen the mole; and of Boreal 
ha^nalit only four or five males have been seen here.'" With most 
of these species (excepting the Tenthredinie) there is at present ni> 
evidence that tbe females are subject to parthenogenesis ; and tlius we 
see how ignorant we are of the causes of the apparent discrepancy in 
the proportion of Ihe two aeies. 

In tba other Claases of tbe Artieukta I have been able to collect still 

" Walsh, in 'The American En- derhBuechreckB,' 1828, p. 20. 

lomologist,' vol. 1. 1869, p. 103. "' Obaei-vations on N. American 

F. Smith, 'Record of Zoological Nenroptera," by H. Hagen and B.D, 

Litsrature,' 1867,5.328. Walsh, ' Proe. Ent. Sot. Phils- 

'• 'Farm Insacts,' pp. 46-46. delphia,' Oct. 1863, pp. 168, 223, 

" 'Anwendung der Darwinachen 239. 

[*hi-e Verh. d. n. V. Jahrg. iiiv.' •• 'Proc. Ent. Soc Londi n,* Kab, 

" 'Die Stricli, Zog oder Wsn- 17, 1861 


RiDip. VllL Proportion OJ iJu Se:cei. 255 

InsB informiitlon. WiU) Spldere, Mr. Blaokwall, who hiis oarefuK/ 
utteiidtd to this olaaa during many yeara, writes to me lluit the maloa 
from their more eimtiu habita are more nommoiily sseii, and thorelbre 
appair more numerous. I'hia Is actually the case with a few species ; 
hut he mentioHfl BBveral species in sii genera, in wtiii'h the females 
appear to be much more nuinemua tlian lie males." The small sizeof 
the males in comparison with the femolej (a peculiarity which is some- 
times carried to an extreme degree), and their widely diBerKat appeaj- 
jnce, may account in some iostanoes for their rarity in oolleetiona." 

Some of the lower Crnstscoans are ahla to propagate their kind 
asesuftlly, and this will actount for tha entieme larity of the males ; 
thus VooSiBboJd** carefully esamined no less than 13,000 specimens of 
Apus from twenty-one localities, and amongst these he found only 
319 males. With some other forms (as Tanais and Cypria), as Fritz 
Midler informs me, there is reason to believe that the males are much 
shorter-lived Hum the females ; and thia would explain their scarcity, 
supposing the two aesea to be at first equal in number. On the other 
hand, Miiller has inyariablj taken far more males than females of the 
Dias^lidie and of Cypridina on the shores of Brazil; thus witli a 
BpGcies in the latter geuns, 63 specimens caught the some day included 
57 males ; but he suggBsts that this preponderance may be due Ic 
some unknown difference in the iiabits of the two seies. With onf 
of the higher Brazilian crabs, numelj a Oelaaimua, Frita MUlIei 
found the males to be more numerous than the feiaales. According 
to the lni;ge experience of Mr. 0. Spence Bate, the reverse seems tc 
be the taso witli six common British crabs, the names of which he 
has given me. 

The proportion qf tM seaes in relation to naltiTat seleclion. 
There is reason to enapeotthat in srane eases man has by 
Bdeetion indirectly influenced his own sex-pioducing powers. 
Certain women, tend to produce dutii^ their whole lives more 
children of one eei than of the other : and tJie same holds good 
of many animals, for instance, cows and horses ; thna Mj". Wright 
of Yeldersley House informs me that one of his Arab mares, 
though put seven times to different horses, produced SBven 
fillies. Though I have very little evidence on this head, analogy 
would lead to the belief, tliat tJie tendency to produce either 
aes would bo inherited like almost every other pecaliaritj, for 
instance, that of. producing twins ; and concerning the above 
tendency a good authority, Mr. J. Downing, has communicated 
to me facts which seem to prove that this does occnr in certain 
fcmiiiea of 8hoj+-horn cattle. CoL Marshall " has recently found 
on careful examination that tlie Toda?, a hill-tribe of ludin. 

•' Another great anthority witi 0. P. Cambridge, u quoted it 

res|«ct to this class, Prof. Thoreli of 'Quarterly Joojnal of Science, 

(Jpsala ('On Enropean Spiders,' 1868, p. 429. 

1869-70, pai't i. p. 205) speaks as if '" Beitiage znr Fartheno^oesii, 

fsinole spidera wore goneially oom- p. 174. 


i$6 The Desc-f>il of Maft Pabt IL 

consist of 112 males and 84 females of all ages — that is in a latdo 
ot 133-3 males to 100 females. The Todas, who are polyandrous 
in their marriages, daring former times invariably practised 
female infanticide ; but this practice has now been discontinued 
for a considerable i)eriod. Of the children boiii withia late years, 
the males are more numerous than the females, in the propoitiou 
of 124 to 100. Colonel Mkraholl accounts for this fact in tJie 
followii^ ii^nlous manner. " Let us for the purpose of illwsti'a- 
" tion take tbree families as representing an average of the 
" entire tribe ; say that one mother gives birth to six daughters 
■' and no sons; a second mother bas sis sons only, whilst tha 
" third mother has tbree sons and tbree daughters. The first 
" mother, following the tribal custom, destroys four daughters 
" and preserves two. The second retains her sis sons. The third 
" kills two daughters and keeps one, as also her three sons. We 
" have then from the three families, nine sons and three daughters, 
" with which to continue the breed. But whilst the males 
" boloi^ to families in which the tendency to produce sons is 
" groat, the females arc of those of a converse inclination. Thus 
" the bias strengthens with each generation, imtil, as we find, 
" families grow to have habitually more sons than daughters." 

That this result would follow from the above form of iufunticido 
seems almost cei-tain ; that is if we assume that a ses-producing 
t-endency is inhoriUd. But as the above numbers are so ex- 
tremely scanty, I have searched for additional evidence, but 
cannot decide whether what I have found is trustworthy ; 
jievertbelees the facts are, perhaps, worth giving. The Maories o£ 
New Zealand have long practised infenticide ; and Mr. Fenton" 
states that he " has met with instances of women who have de- 

"stroyed foar, sis, and even seven children, mostly females. 

' However, the universal testimony of those best qualified to 
" jiii^e, is conclusive that this custom bas for many years been 
" almost estinct. Probably the year 1835 may be named as the 
" period of its ceasing to exist." Now amongst the New Zea- 
landera, as with the Todas, male births arc considerably in excess. 
Mr. Fenton remarks {p. 30), " One fact is certain, although the 
" exact period of the commencement of this singular condition of 
" the disproportion of the sexes cannot be demonstratively fixed, 
" it is quite clear that this course of decrease was in full opiirai- 
" tion during the years 1S30 to 1844, when the nua-adult 
" population of 1844 was being produced, and has continued 
" with great energy up to the present time." The following 
statements are taken from Mr. Fenton (p. 26), but as the nnmbera 

" 'AboHEvaril luhabitiints gf Kew Zealand; Govemmeiit Report,' 1069, 


Cbap, VIII. Proportion of tlte Sex€s. 257 

ate not large, aDd as the census was not accurate, uniform 
results cannot be expected. It Bhould be bome in misd in this 
and the following cases, that the nonnal state of everj popTilation 
is an excess of women, at least in all ciyiliaed countries, chiefly 
owing to the greater mortality of the male sex during youth, and 
partly to accidents of all kinds later in life. In 1858, the 
natiye population of New Zealand was estimated as cousistdi^ 
of 31,667 males and ii4,803 females of ali ages, that is m the 
ratio of ISO'S males to 100 females. But during this same year, 
and in certain limited districts, the numbers were stscertained 
with, much care, and the males of all ages were here 753 
and the females 616; that is in the ratio of 122-2 males to 100 
females. It is more importaut for us that during this same 
yeai of 1858, the non-adalt males within the same district 
were found to be 178, and the -aon-aduit females 142, that is in 
the ratio of 125-3 to 100. It may ba added that in 1844, at 
which period female infantieide had only lately ceased, the 
non-adiiU males in one district were 281, and the non-adidt 
females only 191, that is in the ratio of 144-8 males to 100 females. 

la the Sandwich Islands, the males exceed the females in 
number. Infanticide was formerly practised there to a frightful 
sxtent, but was by no means confined to female infants, as 
is shewn by Mr. Ellis," and as I have been informed by Bishop 
Staley and the Eby. Mr. Coan. Nevertheless, another apparently 
trustworthy writer, Mr. Jarvea," whose observations apply to 
the whole arohipelago, remarks :— " Numbers of women are to 
" be fomid, who confess to the murder of from three to sis or eight 
^ children ,-" and he adds, " females from being consiiiered less 
" useful than males were more often destroyed." From what ia 
known to occur in other x>art3 of the world, this Btaf«ment is 
probable; but must be received with much caution. The 
practice of infanticide ceased about the year 1819, when idolatry 
was abolished and missionaries settled in the Islands. A careful 
census in 1839 of the adult and taxable men and women in the 
island of Kaaai and in one district of Oahu (Jarves, p. 4(M), 
gives 4723 males and 8776 females; that is in the ratio of 
125-08 to 100. At the same time the number ot males under 
fourteen years in Kauai and under eighteen in Oahu was 1797, 
and of females of the same ages 1429 ; and here we have the 
ratio of 125-75 males to 100 females. 

In a census of all the islands in 1850," the males of all ages 

" ' Karrative of s Tnnr ttiougli " Thia is given in tha Rev. H. T, 

Hawaii,' 1826, p. 2B8. Cheever'a 'Lite In itae Sandwich la- 

" ' History of the Saodwiuh lands,' 1851, p. 277. 


Ttie Descent of Man. 

Past II. 

amount to 36^2, and the females to 33.128, or as 109'49 to 
100. The males under seventeen years amounted to 10,773, and 
the females Tmder the same age to S593, or as 112-3 to 100. 
From the census of 1S72, the proportion of males of all ages 
(induding half-castes) to females, is as 12.5-36 to 100. It mast 
be borne in mind that all these returns for the Sandwich 
Islands give the proportion of living males to living females, 
and not of the births ; and jui^ittg from all civilised eountdes 
the proportion of males would have been considerably higher it 
the numbers had referred to births.'" 

From the several foregoing cases we have some reason to 
believe that infanticide practised in the manner above explained, 
tfflids to make a male-producing race ; but 1 am &r from sup- 
posic^ that this practice in the case of man, or some anali^ons 
process with otter species, has been the sole determining cause 
of an excess of males. There may be some unknown law leadh^ 
to this result in decreasing races, which have already become 
somewhat infertile. Besides the several causes previously 

" Dr. Coulter, in describing 

on this anbject from the breediug 

('.'oumal R. Geograph. Soc,' vol. 

of dogs; inasmuch as in most breeds, 

V. 1835, p. 67) the slate of Cali- 

with the eiceptinn, perhaps, of 

fornia about the year 1830, says 

greyhounds, many move female 

tbaC the nativee, reclaimed by the 

puppies are destroyed than males, 

Spimsh missiouBTJes, iiave Dearly 

just as with the Toda infants. Mr. 

all perished, or are perishing, al- 

Cupples assures me that this it 

though well treated, not driven 

usual with Scotch deer-hounds. 

from their nativa land, and kept 

Unfortunately, I know nothing ol 

from the nse of spirits. He at- 

the proportion of the aeies in any 

tributes this, in great part, to the 

DndDDbted last that the men greatly 

there the male births are to the 

«oeed the women in number; but 

female as 110-1 to 100. Now from 

h d t k w wh th th 

quin m d from many breeders, 

d t fell f f m J ff p g 

t eem th i the females are 

t m fmal dy g dur g 

aoro pects more esteemed, 

early y th Th I tt It tl e, 

wrd g t 11 an logy y 

1 d P3 t ppear that the female 

imp b bl H add h t m 

p pt f th best-brei dogs are 

fant d p p ly 11 d .s 

jst m t lly destroyed more than 

!ias bee d m m hed f m h ged 
h bit f 1 f 
I had h ped t ga aom hght 


■ Ckap. Vm. Proportion of tJie Sexes. 259 

alluded to, lie greater facility of parturition anongst BaTagea, 
and the less consequent injury to their male infants, would 
tend to increase the proportion of live-bom males to females. 
There &.<:iv» not, however, seem to be any neceBsary connection 
between savage life and a marked excess of males ; that is if we 
may judge by tlie character of the scanty oflspring of the lately 
existing 'f asmanians and of the crossed ofispring of the Tahitians 
now inhabiting Norf(>lk Island. 

As the males and females of many aninials differ somewhat in 
habits and are exposed in different degrees to danger, it is 
probable that in many cases, more of one sex than of the other 
are habitually deaiioyed. But as far as I can trace out the com- 
f lioatioE of causes, an indiscriminate though large destruction 
of either ses would not tend to modify the sex-producing power 
of the Bpedes. With strictly social animals, such as bees or ants, 
which produce a Tast number of sterile and fertile females in 
comparison witli the males, and to whom this preponderance is 
of paramount importance, ws can see that those communities 
would flourish best which contained females having a strong 
inherited tendency to produce more and more female ; and in 
such cases an unequal sex-producing tendency would be ulti- 
mately gained through natural selection. With animals living 
in herds or troops, in which the males eome to the front and 
defend the herd, as with the bisons of North America and certain 
baboons, it ia coneeiyable that a male-producing tendency might 
be gained by natural selection; for the individuals of the better 
defended herds would leave more numerous descendants. In 
theeaseof mankind the advantage arising from having a pre- 
ponderance of men in the tribe is siipposed to be one chief cause 
of the practice of female infenticide. 

In no case, as fer as we can see, would an inherited tendency 
to produce both sexes in equal nnmbers or to produce one sex 
in excess, be a direct advantage or disadvantage to certain 
individuals more than to others ; for instance, an individual 
with a tendency to produce more males than females would not 
succeed better in the battle for life than an individual with an 
opposite tendency ; and tberefore a tendency of this kind could 
not be gained throi^h natural selection. Nevertheless, there are 
certain animals (for instance, fishes and cirripedes) in which two 
or more males appear to be necessary for the fertilisation of the 
female ; and the maJes accordii^ly largely preponderate, but it 
is by no means obvious how this male-prodncii^ iendency could 
have been aeqmred. I formerly thought that when a tendency 
to produce the two sexes in eqaal numbers was advantageous to 
the species, it wpnld follow from aatural eelection, but I now 


The Descent of Man. 


bEooBDAHY SEST74L Charaotbrb Df THB LowER Classes of 
THK Anisial Kingdom. 

Thesa characters absent in the lowest classes— Brilliant colonrs— Mollnsca 
— Annelids — Crustacea, eecondarf aeinal characters stranglj developed ; 
dimorphism ; colour ; uharacters not acquired before maturitj—Spiders, 
eeiual colours oi'; stridulation by the males — Myiiapoda. 

With animalB beloi^ing to the lower classes, tlie two sexes 
are not rarely united in the same individual, and therefore 
secondary sexual characters cannot he developed. In niany 
cases where the sexes are separate, both are permanently a(> 
tached to some support, nnd the one cannot search or struggle 
for the other. Moreover it is almost certain that these animals 
have too imperfect senses and much too low mental powers, to 
appreciate each other's beauty or other attractions, or to feel 

Hence in these classes or frub-kingdoms, such as the Protozoa, 
Cfelenterata, Eohinodermata, Scolecida, secondary sexual eha- 
ract«rs, of the kind which we have to consider, do not occur; and 
this fact agrees with the belief that such characters in the 
higher classes have been acquired through sexual selection, 
which depends on the will, desire, and choice of either sex. 
Nevertheless some few apparent exceptions occur; thus, as I 
hear from Dr. Baird, the males of certain Entozoa, or intemal 
I)ara3itic worms, differ slightly in colour from the females ; but 
we hare no reason to suppose that such differences have been 
augmented through sexual selection. Contrivances by which the 
male holds the female, and which are indispensable for ths 
propagation of the species, are independent of sexual selection, 
and have been acquired through ordinary selection. 

Many of the lower animals, whether hermaphrodites or with 
separate sexes, are ornamented with the most brilliant tints, or 
are shaded and striped in an elegant manner ; for instance, many 
corals and sea-anemones (Actiniie), some jelly-fish (Medusa, 
Porpita, &c),some Planariie.manystar-flshes, Echini, Ascidians, 
&0. ; but wo may conclude from the reasons already indicated, 
namely the nnion of the two sexes in some of these animals, the 
permanently afSsed condition of othere, and the low mentri 
powers of all, that such colours do not serve as a sexual 
attraction, and have not been acquired through sexual selection. 


Ohat. IX. Sexual i>el4ctton. 261 

It should be harm in mind that in no caso hare we suffi- 
cient evidence that colouis ha^e been thus aeqnited, ei- 
cept where one sex is mnch more hrilljantlj or conspicuously 
colooied than the other, aaid where there ia no difference 
in habits between the seses sufficient to account for their 
different colours. But the evidence is tendered as complete 
as it can ever be, only when the more ornamented indivi- 
duals, almost always the males, voluntarilj display their 
attractionB before the other ses ; for we cannot believe that such 
display is useless, and if it be advantageous, sesua! selection 
will ahnost inevitably follow. Wo may, however, extend this 
conclusion to both sexes, when coloured alike, if their colours are 
plainly analogous to those of one ses alone in certain other 
Epodea of the same greup. 

How, then, are we to account for the beautiful or even 
gorgeous colours of many animals in the lowest classes? It 
appears doubtful whether such colours often serve as a protec- 
tion ; but that we may easily err on this head, will be admitted 
by every one who reads Mr. Wallace's excellent essay on this 
subject. It would not, for instance, at iirst occur to any one 
that the transparency of the Medusas, or jelly-flshea, is of the 
highest service to them as a protection; but when we are 
reminded by Hacfcel that not only tho medusre, but many 
floating mollusca, cmstaceaus, and even small oceanic fishes 
partake of this same glass-like appearance, often accompanied 
by prismatic colours, we can hardly doubt that they thus 
escape the notice of pelagic birds and otter enemies. M. 
Giawi is also convinced' that the br^ht tints of certain 
sponges and ascidians serve as a protection. Conspicuous 
colours are likewise beneficial to many ammals as a warning to 
tkeu would-be devourera that they ai* distasteful, or that they 
possess aome special means of defence ; but this subject will ba 
discussed more conveniently hereafter. 

We can, in oui ^orance of most of the lowest animalB, only 
say that their bright tints result either from the chemical 
nature or the minute structure of their tissues, independently 01 
any benefit thus derived. Hardly any colour is finer than that 
of arterial blood ; but there is no reason to suppcdo that the 
colour of the blood is in itself any advantage; and though it 
adds to the beauty of the maiden's cheek, no one will pretend 
that it has been acquired for this purpose. So again with many 
animals, especially the lower ones, the bile is richly coloured ; 
thus, as I am informed by Mr. Hancock, the extreme beauty of 
the Eohdsa (naked sea-slugs) is chiefly due to the biliary glands 
' 'ArchivsudeZooJog. Eii>^r.,'Oct. 1872, p. 563. 


262 The Descent of Man. Part H. 

teing seen through, tha tranBlucent integuments — tliis beauty 
beii^ probably of no service to these animals. The tints of tiu 
ilocajing leaves in an American tbrest are described by every 
one as gorgeous; yet no one supposes that theee tints are 
of tie least advantage to tie trees. Bearing in mind now many 
subBtances closely analogous to natural oi^anic cowpounda hava 
been receutlj formed by chemists, aad which exhibit the moat 
splendid eolouis, it would hsTe been a strange fa«t if substances 
similarly coloured had not often originated, independently of 
any useful end thus gained, in the complex laboratory of living 

The. svi-Hngdom of t/w AWJwica.— Throughout this great 
division of the animal kingdom, as fer aa I can discover, 
secondary sexual characters, such as we are here considering, 
lever occur. Nor could they be expected in the three lowest 
dasses, namely in the Ascidians, Polyzoa, and Brachiopods 
(constituting the Molluscoida of some authors), for most of 
these animals ore pormanenfiy affixed to a support or have their 
sexes united in the same individual. In the LamelhbrancMata, 
or bivalve shells, hermapkroditiBm is not rare. In the ne.xt 
higher class of the Gasteropoda, or univalve shells, the sexes are 
either united or separate. But in the latter case the males 
never possess special organs for finding, securing, or charming 
the females, or for fighting with other males. As I am informed 
by Mr. Gwyn Jeffireys, the sole external difference between the 
sexes consists in the shell sometimes differing a little in form ; 
for instance, the shell of the male periwinkle {Littorina lUtorea^ 
is narrower and has a mora elongated spire than that of the 
female. But difiereuces of this nature, it may be presumed, are 
directly connected with the act of reproduction, or witt the 
development of the ova. 

The Gasteropoda, though capable of locomotion and furnished 
with imperfect eyes, do not appear to be endowed with sufficient 
mental powers for the members of the same sex to struggle 
ti^ether in rivalry, and thus to aeq^uire secondary sexual 
characters. Nevertheless with the pulmoniferous gasferopods,or 
land-snails, the pairing is preceded by courfstup; for th^e 
animals, though hermaphrodites, ere compelled by their structure 
to pau- together. Agassiz remarks,' " Quicomiue a en I'occasioa 
" d'observer les amours des limagons, no sanrait mettre en dout© 
" la seduction d^ployee dans les mouvements et les allures qui 
" prcpnrent et accomplissent le double embrassement de ces 
" hermaphrodites." These animals appear also susceptible o( 
Bome degree of permanent attachment : an accurate observer, 

' ' Dc TEspJce et de la Class.' &c, 1869, p. 106. 


Chap. tS. Molluscs. 263 

Mr. Lonsdale, informs me that he placed a pair of laad-anailB. 
(^Selix pomatiii), one of which was weakly, into a email and ill- 
proTided garden- After a Bhort time the strong and healtihy 
iudividual disappeared, and was traced h; ita track of slime 
over a wall into an adjoining well-stocked garden. Mr. 
lionsdale concluded that it had deserted its dcMy mate; hut 
after an ahsence of twenty-fom hours it returned, and apparently 
communicated the result of its successful exploration, for both 
then started along the same track and disappeared over the 

Even in the highest class of the Molli^sca, the Cephalopoda or 
cuttlefishes, in which the sexes are separate, secondary sexual 
characters of the present kind do not, as fat as I can discover, 
occur. This fe a surprising circumstance, as these animals 
posses highly-developed sense-organs and have considerabla 
mental powers, as will be admitted by every one who has watched 
their artful endeavours to escape from an enemy.' Certain 
Cephalopoda, however, are characterised by one extraordinary 
sexual character, namely, that the male element collects within 
one of the arms or tentacles, which is then cast off, and clinging 
by its sucking-discs to the female, lives for a time an independent 
life. 80 completely does the cast-off arm resemble a separate 
ajiimal, that it was descritied by Cuvier as a parasitic worm 
under the name of Hectocolyle. But this marvellous structure 
may be classed as a primary rather than as a secondary sexual 

Although with the MoUusca sexual selection does not seem to 
have come into play ; yet many univalve and bivalve shells, 
such as volutes, cones, scalloi>s, il-c, are beautifully coloured 
and shaped. The colours do not appear in most cases to be of 
any use as a protection ; they are probably the direct result, as 
in the lowest classes, of the nature of the tissues; the patterns 
and the sculpture of the shell depending on its manner of 
growth. The amount of light seems to be influential to a certain 
extent; foralthoi^h, as' repeatedly stated by Mr. Gwyn Jeffreys, 
the shells of some species living at a profound depth are brightly 
coloured, yet we generally see the lower surfeices, as weU as the 
parts covered by the mantle, less highly-coloiaed than iho 
upper and exposed surfaces.* In some cases, as with shells 

> See, for instaace, the accoimt influeaee of light on the coIodts d{ 

which I have pven in my 'Jaurnal B froudescent incruBtatioD, d«- 

fjf Researches, 1845, p. 7. pojited hy ike eurf on the coagi- 

' I have given ('Geolog. Obsei- roeks of AseeosLon, and formed by 

rations on Voleanic IsUnds,' 1844, the solution of triturated Bea-sb^U* 
p. 53) a cnriooa instance of the 


264 Tke Dgscmt of Man. piST II. 

living iimoEget corals or brightly-tinted seafweeds, the bright 
colours may servs as a protection.' But that many of the nudi- 
branch mollusca, or sea-sings, are as beautifully coloured as any 
shells, may be seen in Messrs. Alder and Hancocf a magnificeat 
work ; and fi^m information kjndly given me by Mr. Hancock, 
it seems extremely doubtful whether these colours usually serve 
as a protection. With some sj)ecies this may be the ease, fl£ with 
one kind which hves on the greon leaves of algte, and is itself 
brightr^eeu. But many brightly-coloured, white or otherwise 
conspicuous species, do not seek concealment ; whilst again some 
equally conspicuous species, as well as other dul! -coloured kinds, 
live under stones and in diirk recesses. So that with these nudi- 
branch molluscs, colour apparently does not stand in any doso 
relation to the nature of the places which they inhabit. 

These naked sea-slugs are hermaphrodites, yet they pair 
together, as do laod-snails, many of which have extremely 
pretty shells. It is conceivable that two hermaphrodites, 
attracted by each other's greater beauty, might unite and leave 
offspring which would inherit their parents' greater beauty^ 
But with such lowly-organised creatures this is extremely 
improbable. Nor ia it at all obvious how the offepring from the 
more beautiful pairs of hermaphrodites would have any ad- 
voafage over the offspring of the less beautiful, so as to increase 
in number, unless indeed vigour and beauty generally coincided. 
We have not bore the case of a number of males becoming 
mature before the females, with the moro beautiful males 
selected by the more vigorous females. If, indeed, brilliant 
colours were beneficial to a hermaphrodite animal in relation 
to its genera! habits of life, the more brightly-tinted individuals 
would succeed best and would increase in number; but this 
would be a case of natural and not of sexual selection. 

Sab-kingdom 0/ the Vermes: Class, Annelida (or Sea-worms'). — 
In this class, although the sexes, when separate, sometimes 
differ from each other in characters of such importance that they 
have been placed londer distinct genera or even fomilies, yet the 
differences do not seem of the kind which can be safely al- 
iributcd to sexual selection. These animals are often beauti- 
fully coloured, but as the sexes do not differ in this respect, we 
are but little concerned with them. Even the Nemertians, 
though so lowly organised, " vie in beauty and variety of 
" colouring with any other group in the invertebrate series;" yet 

' Dr. Morse has lately disonased ' Proo. Boafoo &ifi. "f Kat. Hist.' 
this Enbjsct in his paper on the vol. liv,, April, 1871, 
AdiiptiTB Coloiatian of Moltnsca, 


Chap, IX. Crmtaceans. 26g 

Dr. Mcintosh" cannot discoi'ui- that these coloUTa are of any 
service. The sedentary annelids become duller-coloured, ac- 
cording to M, Qnatrefagefl,' after the period of reproduction ; and 
this I presume may be attributed to their less Tigorous condition 
at that time. All these worm-like animals apparently stand too 
low m the scale for the indiTidnala of either sex to exert any 
choice in selecting a partner, or for the individuals of the Game 
SOX to struggle together in riTaJry. 

Sah-kingdom of the ATthropoda: Q]aas, CruedKea.— In this great 
class we first meet with nndoubted secondary sexnal characters, 
often developed in a remarkable manner. Unfortunately the 
habits of cmstaceans are very imperfectly known, and we cannot 
explain the uses of many structures pccnliBr to one ees. With the 
lower parasitic speeifis the males are of small size, and they 
alone are furnished with perfect swimming-legs, anlennEe and 
sense-organs; the females being destitute of these organs, with 
their bodies often consisting of a mere distorted mass. But 
these extraordiuary differences between the two seses aro no 
doubt related to *heir widely different habits of life, and eon- 
sequeutly do not concern us. In various crustaceans, belonging to 
distinet families, the anterior antennse are furnished with pecuUar 
thread-like bodies, which are believed to act as smell ing-ot^ns, 
and these are much more numerous in the males than in the 
females. As the males, without any unusual development of 
their olfactory organs, would almost certainly be able sooner or 
later to find the females, the increased number of the sraelling- 
throads has probably been acquired through sexual selection, by 
the better provided males havii^ been the more successful in 
finding partners and in producing ofepring. Fritz Miiller has 
described a remarkable dimorphic species of Tanais, in which the 
male is represented by two distinct forms, which never graduate 
into each otter. In the one form the male is furnished with 
more numerous smelling-threaito, and in the other form with 
more powerful and more eloi^ated chelSB or pincers, which servo 
to hold the female. Frita MUller suggests that these differences 
between the two male forms of the same species may have 
originated in certain individuals having varied in the number of 
the smelling-threads, whOst other individuals varied in the 
shape and size of their cheto ; so that of the former, those which 
were best able to find the female aui of the latter, those which 

• See hia baantifal monograph on See M P rrier, 'I'Origine d< 

'British iiineliJs,' part i. 1873, IH mm d prta Darwin," ' Reiuf 



The Descent of Man. 


were best able to hold her, have left the greatest number of 

progeny to inherit their respective advant^ea." 

In some of the lower crustaceans, the right anterior antenna 

(, of the male differs greatly in strncture 

from the left, the latter resembling in 

ite simple tapering joints the anteimEe 

of the female. In the male the 

modified antenna is either swollen in 

the middle or angularly bent, or 

converted (flg. 4) into an elegant, 

and sometimes wonderfully complex, 

prehensile oj^n." It serves, as I hear 

from Sir J. Lubbock, to hold the 

I female, and for this same purpose one 

' of the two posterior legs (i) on the 

same side of the body is converted 

into a forceps. In another family the 

inferior or posterior antennjo are 

" curiously zigzagged " in the males 


In the higher crustaceans the an- 
terior legs are developed into chela 
or pincers; and these are generally 
laiger in the ma!o than in the female, 
—BO much so that the market value of 
the male edible crab (Cancer pagurus), 
according to Mr. 0. Spence Bate, is 

a. Part of right anwrior ™- ^^° times as great as that of the fe- 

i^» of male, fonniqg a male. In many species the cholte are 

b. pSit«to^pai"of thoracic lep of unequal sizc on the opposite side of 
e nrndrfflmaio *^^ body, the right-hand one being, as 

" ™ °' I am informed by Mr. Bate, gonerally, 

though not invariably, the largest. This inequality is also often 
much greater in the male than in the female. The two ehel» 
of the male often differ in structure (figs. 5, 6, and 7), the 
smaller one resembling that of the fcmala What advantage is 
1 the opposite sides of the 

gained by their inequality i 

' ' Facta and Argumeats for 
Darwin,' EDglisli translat. 1869, p. 
20. See ttie preTious diGcueston on 
the olfactoiy threads. Sars has 
descrihed a somewhat acaJogons 
case (as quoted in ' Nature,' 1870, 
p. 455) in a Norwegian crustacean, 
the Pontoporeia afiiiis. 

' See Sir J. Lubboclc in ' Annals 

and Mag. of Sat. Hist.' vol. li. 
1S53. pi. i. and i. ; and vol. lii. 
(1853) pi. vii. See also Lnbbock in 
'Transact. Ent. Soc' vol. iv. new 
series, 1856-1858, p. 8. With re- 
spect to the zig-zagged antenna 

'Facts I 

i beloH 

z Miller, 


body, and by the inequality being much greater in the male than 
in the female ; and why, when they are of equal size, both are 

of body of dJliunsuBtt (from Milnfi-Edwotds), showing the 
eDtly-couatmclfd rlgUt And left'hajid cbelie of the male, 
lisuiu has revorsed the drawing, ami made tie left-liami cl 

l^g. e. SeoDDd leg i]rma1eOrcbeetia,TticnnClDga (from Fells Milller). 
Rg.7. Mtlo offamale, 

often much larger in the male than in the female, is not known. 
As I hear from Mr. Bate, the chel» are aometiiQes of such length 
and size that they cannot poseibly he used for carrying food to the 
mouth. In the males of certaia fresh- water prawns (PaJtemon) 
the right 1^ ia actually longer than the whole body." The 
grsat size of the one leg with its chete may aid the male in 
fighting with his rivals; but this will not account for their 

"> See a paper by Mr. C. Spence 585. I am greatly indebted to Mr. 

Bate, with figures, in 'Proc.Zoolos. Speoce Bate for nearly alt the aboTa 

Soc.' 1868, p. 363; and on the statements with respect to the chelie 

nomendaturi of the geDus, ibid. p. of the higher cruataceane. 


26S The Descent of Man. Part II. 

inequality in the female on the opposite sides of of the body. In 
GelasiiQus, according to a statement quoted by MilDe-Edwatds," 
the male and the female live in the eamo bnrrow, and this 
shews that they pair ; the male closes the mouth of the burrow 
with one of its ohelie, which is enormously developed ; so that 
hero it indirectly serves as a means of defence. Their main use, 
however, is probably to seize and to secure the female, and this 
in some instances, as with Ganunams, is known to be the case. 
The male of the hermit or soldier crab (Fagurus) for weeks 
together, carries about the shell inhabited by the female." The 
sexes, however, of the common ehore-crab (Vurcimts mana<), s,n 
Mr. Bate informs me, nnite directly after tlie female has moulted 
ber hard shell, when she is so soft that she would be injured if 
seized by the strong pincers of the male ; but as she is caught 
and carried about by the male before moulting, she could then be 
seiKed with impunity. 

Fritz Miiller states that certain species of Melita are distin- 
guished from all other amphipods by the females having " the 
'' coial lamellte of the penultimate pair of feet produced into 
'' hook-Uke processes, of which the males lay hold with the 
" hands of the first pair." The development of these hook-lifco 
processes has probably followed from those females which were 
the most securely held during the act of reproduction, having 
left the largest number of ofllspring. Another Brazilian amphi- 
pod (Orcliesiia Darvdnii, fig. 8) presents a case of dimorphism, 
like that of Tanais ; for there ate two male forms, which differ 
in the structure of their chelte." As either chela would certainly 
BnfSce to hold the female, — for both are now used for this purpose, 
— the two male forme probably originated by some having varied 
in one manner and some in another; both fomis having derived 
certain special, but EearJy! advantages, from their differently 
shaped organs. 

It is not known tlat male crustaceans fight together tor tha 
possession of the females, but it is probably the case ; for with 
most animals when the male is tai^r than the female, be seems 
to owe his greater size to his ancestors having fought 
with other males daring many generations. In most of the 
orders, especially in the highest or the Brachynra, the male is 
larger than the female ; the parasitic genera, however, in which 
the sexes follow difiercnt habits of life, and most of the Ento- 
moBtraca must be excepted. The chele of many crostaeesns are 

" 'Hisl. Nat, dea Crust.' tom. n. of S. Dctoh.' 
1837, p. 50. " Fritz Mtlllei, ' Facts and Argn. 

"Mr. C. Speaca Bate, 'Brit, n-.enls for Darwin,' 1869, pp. 25-38 
Auoo., Fourth Report en Xht Fvuoa 


Chap- IX. 



weapons well adapted for fighting. Thus when a Devil-crab 
(Portiinus paber) was seen by a son of Mr. Bate fighting with a 
Carcmus mcenas, the latter was soon thrown on its back, and had 
eyery limb tern from its body. When several males of a. Brazilian 
Gelaaimns, a species furnished with immense pincers, were 
placed together in a glass veseet by Frita Muller, fhey mutilated 
and killed one another. Mr. Bate put a large male Careinm 


a inte a pan of water, inhabited by a female which was 
paired with a smallei male ; but the latter was soon dispossessed. 


270 The Descent of Man. Part It. 

Mr, Bate adds, " if they fought, tho viotory was a bloodless one, 
■' for I saw BO wounds." This same aatnralist separated a. male 
MDd-skipper (so common on our sea-shores), Gammaras marinut, 
from ite female, both of whom were imprisoned in the same 
vessel with many indiTiduals of the same species. The female, 
when thus divorced, soon joined the others. After a time tho 
male was put again into the same vessel ; and he then, after 
swimming about for a time, dashed into the crowd, and without 
any fighting at once look away his wife. This fact shews that 
in the Amphipoda, an order low in the scale, tho males and 
females recognise each other, and are mutually attached. 

The mental powers of the Crustacea are probably higher than 
at first Eight appeals probable. Any one who tries to catch one 
of the sbore-erabs, so common on tropical coasts, will perceive 
how wary and aJert they are. There is a large crab {Birgm 
Jatro), found on coral islands, which makes a thick bod of the 
picked fibres of the cocoa-nut, at the bottom of a deep burrow. 
It feeds on the (alien, fruit of this tree by tearir^ off the husk, 
fibre by fibre ; and it always begins at that end where the thveo 
eye-like depressions are situated. It then breaks through one of 
these eyes by hammering with its heavy front pincers, and 
turning round, extracts the albuminous core with its narrow 
posterior pincers. But these actions are probably instinctive, so 
that they would ba performed as well by a young animal as by 
an old one. The following case, however, can hardly be so con- 
sidered : a trustworthy naturalist, Mr. Gardner," whilst watching 
a shore-crab (Gelasimns) making its burrow, threw some sheila 
towards the hole. One rolled in, and throe other shells remained 
within a few inches of the month. In about five minutes the 
crab brought out the shell which had fallen in, and carried it 
away to the distance of a foot ; it then saw the three other shells 
lying near, and evidently thinking that they might likewise roll 
in, carried them to the spot where it had laid the first. It 
would, I think, bo difficult to distinguish this act from one 
performed by man by the aid of reason. 

Mr. Bate does not know of any well-marked case of differenco 
of colour in the two seses of our British crustaceans, in which 
respect the sexes of the higher animals so often differ. In some 
cases, however, the males and females differ slightly in tint, bnt 
Mr. Bate thinks not more than may be accounted for by their 
different habits of life, such as by the male wandering more 
about, and being thus more esposed to the light. Dr. Power 

'* "Ttsvels in the Interior of 463, an acoonnt of the habits of the 
BraiiV 1846,p.lll. Itavo giveo, Birgia. 
in m; ' Joanial of Rtseorchei,' p. 


Chjf. is. Crustaceans. 271 

tried to distmgmah by colour the sexee of the Beveral epeciea 
wMch inhabit the Maiiritias, but failed, except with one species 
of Sqnilla, probably S. stylifera, the male of ■which is described as 
being " of a beantiftil blmah-green," with some of the appendages 
chorry-rad, whilst the female is clouded with brown and grey, 
" with the ted about her mnch loss vivid than in the male."" 
In this case, we may suBpect the agency of sexual selection. 
From M. Bert's observations on Daphnia, when placed in a vessel 
illuminated by a prism, we have reason to believe that even the 
lowest crustaceans can distinguish colours. With Saphirina (an 
oceanic genua of Entomostraca), the males are furnished with 
minute shields or cell-hke bodies, which exhibit beautiful 
changing colours ; these ate absent iu the females, and in 
botb sexes of one species." It would, however, be estremely 
rash to conclude that these CTirious organs serve to attract the 
females. I am informed by Fritz Mullet, that in the female of a 
Brazilian species of Gelasiinus, the whole body is of a nearly 
uniform greyish-brown. In the male the posterior part of the 
cephalo-thoras is pure white, with the anterior part of a rich 
green, shadii^ into dark brown ; and it is remarkable that these 
colours are liable to change in the comse of a few minutes— the 
white becoraii^ dirty grey or even black, the greea " losing much 
" of its brilliancy." It deserves especial notice that the males do 
not acquire their br^t colours until they become mature. Thej 
appear to be much more numerous than the females; they 
differ also in the laisei size of their chelte. In some species of 
the genus, probably in all, the seses pair and inhabit the same 
burrow. They are also, as we have seen, highly intell^nt 
animals. From these various considerations it seems probable 
that the male iu this species has become gaily ornamented in 
order to attract or excite the female. 

It has just been stated that the male Gelasunus does not 
acquire his conspicuous colours imtil mature and nearly ready 
to breed. This seems a general mle ia the whole class in respedl 
to the many remarkable structural differences between the sexes. 
We shall hereafter find the same law prevailing throughout the 
great sub-kingdom of the Verfebrata ; and in all cases it is 
eminently distinctive of characters which have been a^uired 
throngh aexnal selection. Fritz Mlillcr''' gives some striking 
instances of this law ; thus the male sand-hopper (Orchestia) 
does not, until nearly full grown, acquire his large claspers, 


•' Mr. Cc. Fraeer, in ' Proc Zoc- 

l.Soi:.'lB69,p,3. I am iodebW 
Mr. Bate for Dr. Fow«'s statc- 

podea,' 18S3, 9. 35. 

" ' Facts and Argnmente,' Ac. 




272 Tli£ Descent of Man. Pabt IL 

which are Terj differently constructed from those of tlie female; 
whilst young, his clitspers resemble those of the female. 

Class, Araclinidd (Spiders). — The seses do not generally diffor 
raacli in coloni, bat the males are often darker than the females, 
na may be seen in Mr. Blackwall's magniSecnt wort." la some 
species, howsTer, the difference is conspicuous ; thas the female 
of SpaTtKsut smaragdiiliis ia dullish green, whilst the adalt male 
has the abdomwi of a fine yellow, with three loi^tudinal stripea 
of rich red. In certain spaoios of Thomisus the sexes closely 
resemble each: other, in othera they differ much ; and analogoas 
cases occar in many other genem. It is often difficult to say 
which of the two sexes depaits most from the ordinary coloration 
of the genus to which the species belong ; but Mr. Blackwall 
thinis that, as a general rule, it is the male ; and Canestriai ■" 
remarks that in certain genera the males can be epecifieally dis- 
tingnished with ease, hut the females with great difficulty. I am 
informed by Mr, Blackwali that the sexes wlulst young usaally 
resemble each other ; and both often undergo great changes in 
colour during their succeesive moults, before airiving at maturity. 
In other cases the male alone appears to change colour. Thus 
the male of the above br^t-coloured Sparassas at first re- 
sembles the female, and acquires his peculiar tints only when 
nearly adult. Spiders are possessed of acute senses, and exhibit 
much intelligence ; as is well kiown, the females often shew 
the. strongest affection for their e^s, which they carry aboat 
enyeloped in a silken web. The males search eagerly for the 
females, and hnve been seen by Canestrini and others to fight for 
poBsesaion of them. This same author says that the union ofthe 
two sexes has been observed ia about twenty species ; and ho 
asserts positively that the female rejects some of the males who 
court her, threatens them with open, mandibles, and at last after 
long hesitation accepts the chosen one. Prom these several 
considerations, we may admit with some eonfideuco tha,t the 
well-marked differences in colour between the sexes of certain 
species are the results of sexaal seleetion ; though we have not 
here the b^t kind of evidence, — the display by the male of his 
ornaments. From the extreme variability of colour in the male 
of some species, for instance of llieridion Uneutwrn, it would 
appear that these sexual characters of the males have not as yet 
become well fixed. Canestrini draws the same conclusion from 

'• 'JL History of Ihe Spiders of 'Caratteri sessnali secondarii de<'l| 

<im,t Britain,' 1861-64. Far the Aiachnidi,' ia the 'Atti della Soa 

("nJlowing facU, see pp. 77, 88, 103. Veueto-Trentina di St Sit. Padora, 

" This author has receotly pub- vol. i. Faac. 'i, 1873, 
liibf^ a vaimble essay on the 


Chap. IX Spiders. 273 

tlie fact tbat the males of certain Gpecies present two forms, 
differing from each other in the size and length of their jaws ; and 
tbis reminds us of the ahore cases of dimorphic crostaceans. 

The male is generally much smaller than the female, sometimea 
to an extraordinary degree,^" and ho is forced to be extremely 
cautioas in making his advances, as the female often carries her 
coyness to a dangerous pitch. De Geet eaw a male that " in tho 
" midst of his preparatory caresses was seized by the object of 
" his attentions, enveloped by her in a web and then devoured, a 
" sight which, as he adds, filled him with horror and indignation." " 
The Kev, 0. P. Cambridge"" accounts in the following manner 
for the extxeme smallness of the male in the genus Nephila. 
" M. Vinson gives a graphic acconnt of the agile way in which 
" the diminutive male escapes from the ferocity of the female, by 
" gliding about and playing hide and seek over her body and 
" along her gigantic limbs : in sneh a puisoit it is evident that 
" the chances of escape would be in fevour of the smallest males, 
" while the lai^r ones would fall early victims ; thus gradually 
" a dinjiuntive race of males would be selected, until at last they 
" would dwindle to the smallest possible size compatible with the 
" exercise of their generative functions, — in fact probably to the 
" size we now see them, i.e., so small as to be a sort of parasite 
'• upon the female, and either beneath her notice, or too agile and 
'■" too small for her to catch without great diflieulty." 

Westring has made the interesting discovery that the males 
of several species of Theridion'^ have the power of malting 
a fitriduhiting sound, whilst the females are mute. The ap- 
paratus consists of a serrated ridge at the base of the abdomen, 
against which the hard hinder part of the thorax is rubbed ; and 
of this stractujre not a trace can be detected in the females. It 
deserves notice that several writere, including the well-known 
arachnologist Walokenaer, have declared that spiders are attracted 
by music." From the analogy of the OrthopteraandHomoptera, 

» Aug. Vinson {' Arsn^das des tion to EntginnlDBy,' vol. I. 1818, 

lies de la Rennion,' pi. vi. tigs. I p. 280. 

and 2) gives a good instance of the " ■Proc.Zoolog.Soc' 1871,p.6ai, 

small idze of the male, in Epeira " 'fheridiiai [Asagma, Sond.) 

nigra. In this species, as I maj lerratipes, ^-pimdatam et gutta- 

add, the male is testaceous and the lam; see Weslring, in Eioyer, 

female black with lega banded with * Naturbist. Tidakrift,' vol. ir. 1843- 

red. Other even more striking 1843, p. 349} and voL ii. 1846- 

eases of inequality in aii» between 1B49, p. 342. See, also, for othei 

the seisi have been recoiiled species, ' Araneai Snecicie,' p. 184, 
('Quarterly Journal of Science," » Dr. H. H. van Zoutevaen, la 

18*8, July, p. 429); bnt I have his Ontch translation of this •ork 

lot seen the original aoconnts. (vol. i, p. 444), has t.illei:f*J. aevanJ 

" Kirhj and Spencc, ' lutrodne- casoi. 


2/4 "^l^ Descent of Man, Part IL 

to be described in the next chapter, we may feel almost sure that 
the stridulatiOD serves, as Westring also belieYes, to call or to 
excite the female; and this is the first case known to me in the 
ascending scale of the animal kingdom of sounds emitted for 
this purpose.'" 

Claasj Myria-poda. — In neither of the two orders in this class, 
the millipedes and centipedes, can I find any well-matkea 
instances of such eesua) difierences as more particularly concern 
ns. In Qlomttris limiata, however, and perhaps in some few 
other species, the males differ slightly in colour from the females ; 
but this Glomeris is a highly variable species. In the males of 
the Diplopoda, the legs belonging either to one of the anterior or 
of the ixisterior segments of the body are modified into pre- 
hensile hooks which serre to secure the female. In some species 
of lulns the tarsi of the male are furnished with membranous 
snekers for the same purpose. As we shall see when we treat 
of Inseclfi, it is a much more unusual circumstance, that it is 
the female in Liihobius, which is furnished with prehensile 
;s at the eitremity of her body for holding the male.* 

Sboondaby SastTAL Chaeacibks or Insects. 

DJTersified structures posaeaa&l by the males ftff seizing the females — 
BifFereacefi between the sciea, ef which the zneaciag ia net understood — • 
Difference in size between the seies — Thysannra — Diptera— Hemiptera 
— Homoptera, miiaical powers posseesed by the malea alone — Orthoptera, 
musical instraments of the males, modi diversified ia Etroclnre ; 
pngnacitj; colours — Kenroptera, seiual differences in colour — Hyme- 
noptera, pugnacity and coloui*— Coleoptera, colonrs; ftirnished with 
great horns, apparently as an ornament; latties; stridnlating organs 

In the immense class of jasocts the sesos sometimes differ in 
their locomotive-organs, and often in their sense-oi^ans, as in 
the pectinated and beautifully plumose antemtffl of the males of 
many species. In Chloeon, one of the EphemeHB, the male has 
great pillared eyes, of which the female ia entirely destitute." 
The ocelli are absent in the females of certain insects, as in the 

" Hilgend(rf, however, has lately 'Hist. Nat. dea Inseitea : .\pteres,* 
ttalleil attention to an analogous torn. it. 1847, pp. 17. IS, k8. 
ttrncture in some of the higher ' Sir J. Lubbock, 'Transact. 

to prodnce sound; Me 'Zoological 484. With respect to the Mu- 

Becord,' 1869, p. 603. tillil» see Westwood, ' Modern 

» Wi^lckenaer et P. Gerrais, Claw, of IiHett<,'Tol. ii. p. 213. 


CEip. X. Insects. 275 

Mutiliidie; and hero tlio females aid likewise wigless. But 
W6 are chiefly concerned with stractuies by which one male is 
enabled to conquer another, either in battle or conrtBhip,thiongh 
his strength, pugnacity, ornaments, or music. The innnmerahle 
contiiTances, therefore, by which the male is able to Eeizo the 
female, may be briefly paesed over. Besides the complex straetures 
at the apex of the abdomen, which ought perhaps to be ranked 
as primary organs,^ " it is asfeMiiabing," as Mr. B. D. Walsh* has 
remarked, " how many different organs are worked in by nature 
" for the seemingly insignificant object of enabling the male to 
" grasp the female firmly." The mandibles or jaws are some- 

. times used for this purpose ; thna the male CorydaUs cornutm (a 
neuropterousinseet in some degree allied to the Dragon-flies, &c) 
has immense curved jaws, many times longer than those of the 
female ; and they are smooth instead of being toothed, so that 

. he is thus enabled to seize her without injury.' One of the 
stag-beetles of North America (J^tuxinus elaphus) uses Ms jaws, 
which are much larger than those of the female, for the same 
purpose, but probably likewise for fighting. lo one of the 
sand-wasps (AmriiopMla) the jaws in the two sexes are closely 
ahTie, buji are used for widely different purposes : the males, as 
Professor Westwood observes, "are exceedingly ardent, seizii^ 
" their partners round the neck with their Bickle-sbaped jaws;'" 
whilst the females use these organs for burrowing in saud-bauks 
and making their nest& 

The tarsi of the fiiant-Iogs are dilated in many male beetles, or 
are furnished with broad cushions of hairs ; and in many genera 
of water-beetiea they are armed with a round flat sucker, so that 
the male may adhere to the slippery body of the female. It is a 

' These organs in the male often species haring been olMerved in 

diSei in cloself-allied species, and oaian. Mr. MaeLachlan informs 

afford eicellent speeifio characters, me (Tide ' Stett. Ent. Zeitnng,' 

But tlieir importance, from a fnn^- 1S67, s. 155) that when several 

tional point of view, as Mr.' R. species of Phryganidtt, whicli pr«- 

MacLachlan haa remarked to me, fieut stron^ly-pronoanced difference* 

has probably been overrated. It of this kiad, were confined together 

nas besD euggested, that siigfat dif- bj^ Dr. Aug. Meyer, they couptedy 

ferencea in these organs would and dds pair produced fertile ova. 
Bullice to prevent the inUrcrossing ' ' The Practical Entomologist,' 

ofwell-markedrarietiesorincipient Philadelpliia, vol. ii. May, 1S6T, 

•peclei, and would thns aid in their p. 88. 

development. That thie can hardly ' Mr. Walah, ibid. p. 107. 
be the ease, we may infer iVom tbe ' 'Modem classijicaticn of la- 
many recorded cases (see, for in- sects,' vol. ii. 1840, pp. 205, 206. 
instance, Bronn, ' Gescbicbte dei Mr. Walsh, who called my attention 
Xator,' b. ii. 1S43, a. 164{ and to the double use of the jaws, say* 
Wwtwcod, ' Transact. Eat. Soc.' that he has repeatedly obearrsj 
RiL iiL IB42, p. 195) of dutinct this fact. 



The Descent of Man. 

mucli more unusual ciicumstance that the female of some water- 
beetles (Djtiscus) have their elytra deeply groored, and in 
Acilias BiiicaiMs tliickly set with Iiairs, as an aid to tbo male. 
The females of some other water- 
beetles (Hydioporus) have their 
elytra punctured for the same 
purpose.* In the male of Crahro 
cribrarim (fig. 9), it IB the tibia 
which is dilated into a broad 
homy plate, with minut« mem- 
braceous dots, giving to it a sin- 
gular appearance lite that of a 
riddle.' In the male of Penthe 
(a genua of beetles) a few of the 
middle joints of the antennje are 
dilated and furnished on the in- 
ferior surface with cushions of hair, 
exactly like those on the tarsi of 
the Carabidte, "and obviously for 
" the same end." In male dragon- 
flics, "the appendages at the tip 
" of the tail sre modified in an 
" almost infinite variety of curious 
" patterns to enable them to em- 
' " brace the neck of the female." 
Lastly, in the males of many in- 
eecls, the legs are famished with peculiar spines, knobs or 
spurs ; or the whole 1^ is bowed or thickened, but this is by no 
means invariably a sexual character; or one pair, or all three 
pairs are elongated, sometimes to an extravagant length.^ 

The sexes of many species in all the orders present differences, 
of which the menning is not understood. One curious case is 
that of a beetle (flg. 10), the male of which has the left mandible 
much enlarged; bo that the mouth is greatly distorted. In 
another Carabidous beetle, Eurygnathus," we have tbo case. 

<• We hav, 
eiplicable case of dimorphism, for 
me of the females of four Euro- 
of Dytiscus, and of 
™riaiQ species of Hydroporus, have 
their elytra smooth; and no inter- 
mediate gradatioDS between the 
Bnlested or punctured, and the quite 
smooth elytra hare beea obserced. 
S«e Dr. H. Schaum, as quoted in 
1 ■ Zoologist,' vol. T.-vi. 1847-48, 

■ Introductioi 


lology,' vol. 

' Westwood, 'Modern Class,' vol. 
ii. p. 193. The following state- 
ment about Penthe, and others in 

My. Walsh, 'Practical Entomolo- 
gist,' Philadelphia, vol. ii. p. B8. 

' Kirby and Spence, ' Introduot." 
Sic., vol. iii. pp. 332-336. 

• ' Insecti Madcrensia,' 1854, p. 


Chap. X. 



tmic[tte as far OS known to Mr. Woliaston, of the head of the 
female being much broader and lai^er, though in a Tariable 
degree, than that of the male. Any mimber 
of such cases could be given. They abound 
in the Lepidoptera: one of the most extra- ' 
ordinary is that certain male butterflies 
have their fore-le^s more or less atrophied, 
with the tihite and tarsi reduced to mere ru- 
dimentary knobs. The wings, also, in the two 
seses often differ in neuration," and some- 
times considerably in outlme, as in the AH- 
con's epitus, which was shewn to me in the 
Btitiah Museum by Mr. A. Butler. The males 
of certain South American butterflies have 
tufts of hair on the margins of the wings, 
and horny excrescences on the discs of the 
posterior i)air." In several British butter- 
flies, as shewn by Mr. Wonfor, the males alone 
are in parts clothed with peculiar scales. 

The use of the bright light of the female ' 
glow-worm has been subject to much discus- 
sion. The male is feebly luminous, as are the 
larvsB a!id even the eggs. It has been sup- 
i by some authors that the light serves to 
away enemies, and by others to 
guide the male to the female. At last, Mr. 
Belt^ appears to have solved the difficulty: 
he iinds that aii the Lampyridae which he haa 
tried are highly distasteful to insectivorous 
mammals and birds. Hence it is in accordance 
with Mr. Bates' view, hereafter to be explained, 
that many insects mimic the Lampyridse f-« ' ? 

closely, in order to he mistaken for them, and pi^^ u,^ TapteoderoB 
thus to escape destruction. He further be- dlatoVtug mncii ™- 
lieves that the luminous species profit by »r^njaie f towM 
being at once recognised as unpalatable. *Kure, mm^e. 
It is probable that the same explanation may be exteaded to the 

'• E. Doubleday, ' Annala and 74. Mr. Wonfor's obserrations are 

Mag. of Nat. Hist.' vol. i. 1848, p. qaofed in ' Popnlar Science Review,' 

379. I nia,y add that the wiags in 1868, p. 343. 

certain Hymenoptera (see Shuckard, " ''ITie Naturalist in Nicaragna,' 

'Foasorial Hymeaop." 1837, pp. 39- 1874, pp. 316-320. On the phos- 

43) differ ia neum^Dii according to plioresctnce of the eg^,see 'Annals 

sei. and Mag. of Nat. Hist.' 1871,' Nov., 


37^ 7ke Descent of Man, Fisr I]. 

Elftterg, both sexes of wHcli are liigUj iuminous. It is not 
kaown why the irings of the female glow-worm have not been 
developed ; but in her present state sho closely resembles a, 
larva, and as larvaa aj^e so lately preyed on by many animals, 
we can understand why she has been rendered bo much mora 
luminous and conspieuous than the male ; and why the larvae 
themselves are likewise luminoas. 

Difference in. Size Txtween ike Sexes, — With insects of all kinds 
the males are commonly smaller than the females; and this 
difference can often be detected even in the larval state. So 
considerable is the difference between the mate and female 
cocoons of the silk-moth (^Bomiyx mori), that in Prance they are 
separated by a particular mode of weighing." In the lower 
classes of the animal kingdom, the greater size of the females 
seems generally to depend on their developing an enonaons 
number of ova ; and this may to a certain extent hold good with 
insects. But Dr. Wallace h^ suggested a much more probable 
srplanation. He finds, after carefully attending to the develop- 
ment of the caterpillars of BonQiyx cynthia and yamamai, and 
especially to that of some dwarfed caterpillars reared from a 
second brood oa unnatnral food, " that in proportion as the in- 
" dividual motli is finer, so is the time required for it? metamor- 
"phosis loiter; and for this reason the female, which is the 
" larger and heavier insect, from having to carry her numerous 
" eggs, will be preceded by the male, which is smaller and has 
" less to mature." " Now as most insects are short-lived, and as 
they are exposed to many dangers, it would manifestly be ad- 
vantageous to the female to be impregnated as soon as possible. 
This end would be gained by the males beii^ first matured in 
large numbers ready for the advent of the females ; and this 
again would naturally follow, as Mr. A. E, Wallace has re- 
marked," through natural selection; for the smaller males 
would bo first matured, and thus would procreate a largo 
number of offspring which would inherit the reduced size of 
their male parents, whilst tho larger males from being matured 
later would leave fewer oSspring. 

There are, however, exceptions to the rule of male msects 
being smaller than the females : and some of these exceptions are 
intelligible. Size aud strength would be an advantage to the 
males, which fight for the possession of the females; and in 
these cases, as with the stag-beetle (Lucanus), the males are 
larger than the females. There are, however, other beetles 

" Eoblnet, 'Ven & Soie,* 1848, sol. v. p. 486. 
P- 207. " ' JoUTDal of Pioc. Enl, Set, 

" 'TrsBMct.Ent, Soc'Srd series, Feb. 4lb, 1867, p. Isii, 


Chap. S, Thysanura. ^79 

which are not known to flght together, of which the males 
exceed the females in size ; and the meaning of tliiB fa«t is not 
known ; but iu some of tlieBe cases, as with the biigo Djnastes 
and Megasoma, we can at; least see that there would be no 
necessity for the males to be smaller than the females, in order 
to be matured before them, for these beetles are not sbort-liTed, 
and there would be ample time for the pairing of the sexes. 80 
again, male dragon-flies (Libellulid^) are sometimes sensibly 
larger, and nerer smaller, than the females ; " and as Mr. 
MaeLachlan believes, they do not generally pair with tho females 
nnta a week or fortnight has elapsed, and imtil they have 
assumed their proper masculine colours. But the most curious 
case, shewing on what complex and easily-overlooked relatioos, 
so trifling a cbaracter as difference in size between the sexes 
may depend, is that of the aculeate Hymenoptera ; for Mr, F, 
Smith informs me that throughout nearly the whole of this 
large group, the males, in accordance with the general rule, are 
smaller than the females, and emerge about a week before them ; 
bnt amoi^t the Bees, the males of A'pii ■mdlifioa, Anthidimn 
manicatum, and Anthopkora acervorwn, and amongst the Fossorea, 
the males of the Methoca ichneumonides, are larger than the 
females. The explanation of this anomaly js that a marriage 
flight is absolutely necessary with these species, and tho male 
requires great strength and size in order to carry the female 
through the air. Increased size has here been acquired in op- 
position to the usual relation between size and the period of 
development, for the males, though larger, emerge before the 
smaller females. 

We will now review the several Orders, selecting such facta 
as more particularly concern us. The Lepidoptera (Butterflies 
and Moths) will be retained for a separate chapter. ■ 

Order, Thymnura.—Tae members of this lowly organiBetl 
order are wingless, dull-coloured, minute insects, with ugly, 
almost misshapen heads and bodies. Their sexes do not differ 
bnt they aie interesting as shewing us that the males pay 
sedulous court to the females even low down in the animal 
scale. Sir J. Lnbboek " says : " it is very amusing to see these 
" littlo creatures (Smj/nthurus luteus) coquetting together. The 
" male, which is much smaller than the female, runs round her, 
" and they butt one another, standii^ face to face and moving 

on the size of the Mies, see Kirby " 'Transact. Limienn Sw." toL 

Md Spenee, ibid. vol. iii. p. 300 ; jiiYi. 1B63, p, 296. 
VD the duratioB of life in insects, 


2So The Descent of Man. PaktIL 

" backn^ard and forward like two playfnl lambs. Then tha 
" female pretends to run awaj and the male runs after her with 
" a queer appearance of anger, gets in front and stands facing 
" her again ; then she turns coyly round, but he, quicker and 
" raoro active, scuttles round too, and seems to whip her 
" with his antennse; then for a bit they stand face to face, 
" play with their antennie, and seem 1« be all in all to one 
" another." 

Oi-der, Vi'ptera (Flies).— The sexes differ httle in colour. The 
greatest difference, known to Mr, P. Walker, is in the genus 
Bibio, in which the maJes are blackish or quite biack, and the 
females obscure brownisli-orange. The genus Elaphomyia, dis- 
covered by Mr. Wallace" in New Giiinen., is highly remarkable, 
as the males are furnished with horns, of which the females are 
quite destitute. The horns spring from beneath the eyes, and 
curiously resemble those of a stag, being either branched or pal- 
mated. In one of the species, they equal the whole body in 
length. They might be thought to be adapted for fighting, but 
as in one species they are of a beautiful pink colour, ei^ed with 
black, with a pale central stripe, and as these insects have 
altogether a very elegant appearance, it is perhaps more probable 
that they serve as ornaments. That the males of some Diptera 
tight together is certain; for Prof. Westwood" has several times 
seen this with the Tipulte. The males of other Diptem ap- 
parently try to win the females by their music; H. MtiUer" 
watched for some time two males of an Bristalis courting a 
female; they holered above her, and flew from side to side, 
making a high buimning noise at the same time. Gnats and 
mosquitoes (Culicidaj also seem to attract each other by hum- 
ming; and Prof. Mayer has recently ascertained that the hairs 
On the antennfe of the male vibrate in unison with the notes of a 
tuning-fork, within the range of the sounds emitted by the female. 
The longer hairs vibrate sympathetically with the graver notes, 
and the shorter hairs with the higher ones. Landois also asserts 
that he Las repeatedly drawn down a whole swarm of gnats by 
uttering a particular note. It may be added that the mental 
fjoulties of the IMptera are probably higher than in most other in- 
sects, in accordance with their highly developed nervous system,*' 

" ' The Malay Archipelago,' vol. " See Mr. B. T. Lowne'a intereet- 

ii, 1S69, p. 313. . ing wQrt, ' On the Anatoiny of the 

" 'Modern Classification of In- Blow-fly, Mnsca yomitoria,' 1870,p. 

leets,' Tol. ii. 18*0, p. 526. 14. He remarks (p. 33) that, " the 

™ Anwendang, &c., 'Verh. d. n. " captured flies utter a peculiar 

?. Jaiirg.' nil. p. 80. Mayer, iu " plaintive note, and that this sauE4 

' American Maturaliat,' 1874, p. 836. " causes othei' flies to disappear." 


Chap. X. Hemiptera and Hoinoptera, 281 

Order, Eemi'pUra (Field-Bugg),— Mr. J. W. Donglas, who has 
particularlj attended to the British species, has kindly given me 
an accoimt of their sexnal differences. The males of soiae spedes 
are furnished with wings, whilst the females are wingless ; tha 
soxes differ in the form of their bodies, elytra, antennsB and tarsi ; 
but as the s^nilicAtion of these differences are unknown, they 
may be here passed over. The females are generally larger and 
more robust than the males. With Britisb, and, as far as 
Mr. Douglas knows, with exotic species, the seses do not 
commonly differ much in colour ; hut ia about sis British 
species the male is considerably darter than the female, and 
in about four other species the female is darker than the male. 
Both sexes of some species are beautifully coloured; and as 
these insects emit an extremely nauseous odonr, their con- 
spicuous colours may serve as a signal that they are nnpalat- 
able to insectivorous animals. In some few cases their colours 
appear to be directly protective : thus Prof. Hoffmann informs 
me that he could hardly distinguish a small pink and green 
species from the buds on the tiunis of lime-trees, which this 
insect frequents. 

Some species of Kednviiiffi make a stridulating noise; and, in 
the ease of Pirates slrididus, this is said''^ to be effected by the 
movement of the neck within the pro-thoracic cavity. Accord- 
ing to Westiing, Beduviae peraonatus also stridulates. But I 
have no reason to suppose that this is a sexual character, ex- 
cepting that with non-social insects there seems to be no use 
fbr sound- producir^ oceans, unless it be as a sexual call 

Order, Homoptera, — Every one who has wandered in a tropi- 
cal forest must have been astonished at the din made by ttio 
male Cicadse. The females are mute; as the Grociaa poet 
Xenarchns says, *' Happy the Cicadas livo, since they all have 
" voiceless wives." The noise thus made could be plainly heard 
on board the " Beagle," when anchored at a quarter of a mile 
from the shore of Brazil ; and Captain Hancock says it can be 
heard at the distance of a mile. The Greeks formerly kept, and 
the Chinese now keep theso insects in cages for tho sate of 
their song, so that it must be pleasing to the ears of some men," 
The Cicadidm usually sing during the day, whilst the rnlgorida 
appear to be night-songsters. The soraid, according lo Landois,'* 

" Westwood, 'MoUem Class, of also, on the Fulgoridffi, Kirbj end 

Inaeits,' vol. ii. p. 473. Speoce, ' Introduct.' vol. ii. p. 401. 

" These particulars ore taken " 'Zeitsehrifl fdr wisseusctaft 

&™ii Weatwooii'a ' Modem Claas. of Zoolog.' B. jvii. 1B67, ». 158-158. 
InseoM," vol. ii 1840, j.. *22. See, 


282 The Descent of Man. PabtH 

is produced by the vibration ot the lips of tte spintcleB, which 
are set into motioii by a current of air emitted from the tracbew; 
but this view has lately been disputed. Dr. Powell appears to 
have proved '^ that it is produced by the vibration of a mem- 
brane, set into action by a special muscle. In the living insect, 
whilst stndulatmg, this membrane can be seen io vibrate ; and 
m the dead insect the proper sound ia heard, if the muscle, 
when a little dried and hardened, ia pulled with the point of a 
pm In the female the whole complex musical apparatus is 
present, but is much less developed than in the male, and is 
never used for producing sound. 

With respect to the object of the music, Dr. Eartman, ia 
spcakiiig of the Cicada, septemdecim of the United States, says, " 
" the drums are now (June 6th aud 7th, 1851) heard in all 
" directions. This I believe to be the marital summons from 
" the males. Standing in thick chestnut sprouts about as high 
" as my head, where hundreds were around me, I observed the 
" females ooming around the drumming males." He adds, " this 
" eeaaon (Aug- 1868) a dwarf pear-tree in my garden produced 
" about fifty larvse of Cic. pniinosa ; and J several times noticed 
" the females to alight near a male while he was uttering hia 
" clanging notes." Jrita Miiller writes to me from S. Brazil 
that he has often listened to a musical contest between two or 
three males of a species with a particularly loud voice, seated 
at a considerable distance from each other : as soon as one had 
finished his song, another immediately began, aud then another. 
As there is so much rivalry between the males, it is probablo 
that the females not only find them by their sounds, but that, 
like female birds, they are excited or allured by the male with 
the most attractive voice. 

I have not heard of any well-marked cases of omnmental 
differences between the seses of the Homoptora. Mr. Douglas 
infonuB me that there are three British species, in which the 
male is black or marked with black bands, whilst the females are 
pale-coloured or obscure. 

Order, Orthopiera (Crickets and GrasBhoppers). — The males in 
the three saltatorial femilies in this Order aie remarkable for 
tttoi TOUsical powers, namely the Achetid» ot crickets, the 
liOcnstidsB for which there is no equivalent English name, and the 
Acridiidro or grasshoppers. The Btridulation produced by some 

" 'TranBact. New Zealand Id- 
•lilnte,' vol. v. 1873, p. 386. 

« I »m indebted to Mr, WalA 




of the Looufitidfe is so loud that it can be heard during the night 
at the distance of a mile ; '" and tliat made by certain species k 
not Timmisical even to the hianan eai, so that tJie Indians on tlie 
Amaaons keep them in wicker cages. AU. observers agree that 
the sounds serve either to call or excite the mute females. "With 
respect to the migratory locusts of Russia, Korte has given ^ an 
interesting case of selection by the female of a male. The males 
of this species (Parihytylus migratorius) whilst coupled witfc the 
female stridulate from ar^er or jealousy, if approached by otlier 
males. The house-cricket when surprised at night uses its -voice 
to warn its fellows,'* la North America the Katy-did {Platy- 
phyllam concavum,oQ6et Hie Locustidie) is described '^ as mount- 
ing on the upper branches of a tree, and in the evenii^ beginning 
" his noisy babble, while rival notes issue from the neighbouring 
" trees, and the groves resound with the call of Katy-did-she-did 
" the live-long night." Mr. 
Bates, in speaking of the Euro- 
pean fieldnsricket (one of the 
Achetidie), says, " the male has 
been observed to place him- 
self in the evening at the 
entrance of his burrow, and 
stridulate until a female ap- 
proaches, when the louder 
notes are succeeded by a 
more subdued tone, wijilst 
the successful musician ca^ 
resses with his antennte the 
roate he has won."'' Dr. 
Scudder was able to excite one 
of these iiwecta to answer him, 
by rubbing on a file with a 
quill.*^ In both sexes a re- 
markable auditory apparatus 
has been discovered by Von Siebold, situated in the front legs. 

rig.ll. GrjUnseiBi 

ipestiia (from LuidolB). 

Left-lumd fl^re, npper Burfdce of vj\rt^ 
cover, wilt IhB projf aSng. Bmnoa mtvan. 
T, acroaa which the teetS (a(j sre soriped. 

" L, Guilding, ' Transact Linn. 
Soc' voL IV. p. 154. 

*' I state this on the authority 
of Koppen, ' Ueber die Heusehrecken 
in Sildrnssland,' 1866, p. 32, for I 
have in vain endeavonred to procure 
Korte's wort. 

=' Gilliert WMte, 'Hat. Hist, of 
SellxJTne.' vol. ii. 1825, p. 262. 

" Harris, 'Insects of New Eng- 
land,' 1842, p. • "" 

•The ^ 

a the 

zons," vol. i. 1863, p. 252. Mr. 
Bates gives a veiy interesting dis- 
cussion on the gmdaldoca in the 
muacal apparatus of the thre9 
families. See also Westwood, 
'Modern Class.' vol. ii. pp. 445 

" ' Proo. Boston Sob. of Nat. 
Hist.' vol, Ii. April, 1868. 

" ' Nouvean Manuel d'Anat. 
Cotnp.' 'French tranalat.), tom. 1 
1850, p. 5S7. 


284 T^Ii-e Descent of Man. Part IL 

In the three Families the sounds are diiTerently produced. In 
iho males of the Achetidio both wing-covers hate tho ssme 
apparatus ; and this in the field-cricket (GryUus campesiris, 
fig. 11) consists, as described by Landoie,'* of from 131 to 138 
sharp, transverse ridges or teeth (s() on the under side of one of 
the nervures of the wing-cover. This toothed nervure is rapidly 
scraped across a projecting, smooth, hard ner- 
Ture (r) on the upper surlace of the opposite 
wing. First one wing Js robbed over the 
other, and then the movement is reversed. 
Both wings are raised a little at the same 
time, so as to increase the resonance. In 
some species the wing-covers of the males are 
furnished at the base with a talc-like plate.^ 
I here give a drawii^ (fig. 12) of tiie teeth on 
the tinder side of the nervnre of another 
Fig, 13. Teetk of Ncr- speeies of Gryllus, Tiz., G. domesticus, "With 
Ss'ffrimiluifd'oTsjr respect to the formation of these teeth, Er. 
Gmber has shewn "" that they have been de- 
veloped by the aid of selection, from the minute scales and hairs 
with which the wings and body are covered, and I cams to the 
same conclusion with respect to those of the CJoleoptera. But 
Dr. Gruber further shews that their, development is in part 
directly due to the stimulus from the friction of one wing over 
the other. 

In the Locustidje the opposite wing-covers differ from each 
other in sfruotm^ (flg. IS), and the action cannot, as in the 
last family, be reversed. The left wing, which a«ts at the 
bow, lies over the right wing which serves as the fiddle. One 
of the nervures («) on the under surface of the former is 
finely serrated, and is scraped across the prominent nervures 
on the upper surface of the opposite or right wing. In our 
British i'hasgonara viridissimii it appeared to roe that the 
serrated nervure is rubbed against the rounded hind-comer 
of tho opposite wing, the edge of which is thickened, coloured 
brown, and very sharp. In the right wing, but not in the left, 
there is a little plate, as transparent as talc, surrounded by 
nervures, and caJled the speculum. In Ephippiger viHum, a 
member of this same family, we have a eurions suhordinato 
modification; for the wing-covers are greatly reduced in size, 
but " the posterior part of the pro-thoras is elevated into a kind 

" ' Zeif Bchi'ift Pir wissfnsehaft. " 'Ocber der Tonapparat der 

Zoolof;.' B. iTii. 1867, s. 117. locnBtiden, ein Beit.-ag zum Dar- 

» Westwood, 'Modem Class, of wiaismus,' 'Zeitsch. fur wissensch. 

Insects,' vol. 1. p. 440. Zoolo^.' B. ssii. 1872, p. 100. 



" of dome over the wing-covers, and whicli has probably the 
" efleet of increasing the soand." "^ 

Me- 13. CblQTociElue To 

We thaa see that the musical apparatus is more differentiated 
or specialised in the Loeustidse (which include, I believe, the 
most powerful performers in the Order), than in tlie Achetidse, 
in which both wing-covers have the Bame structure and the 
same fnnction." Landois, however, detected in one of the 
Locuatidte, namely in Decticus, a short and narrow row of small 
teeth, mere rudiments, on the inferior surface of the right wing- 
cover, which underlies the other and is never used as the bow, 
I olKerved the same rudimentary structure on the under side of 
the right wing-cover in I'liasymiura viridissima. Hence we may 
infer with confidence that the LocRstidte are descended from a 
form, in which, as in the existing Achetidffi, both wing-covers 
Siad serrated nerVures on the under surface, and could be 
, indifferently used as the bow; but that in the Locustidte the 
two wing-covers gradually became differentiated and perfected. 



The Descent of Man. 


on the principle of the division of iabom-, the one to act ex- 
clusivelj as the bow, and the other as tho fiddle. Dr. Gruber 
takes the same view, and has shewn that nidimeatary teeth are 
commonly found on the inferior surface of the right wing. By 
what steps the more simple apparatus in the Achetidje originated, 
we do not know, but it is probable that the basal portions of 
the wing-covers origically overlapped each other as they do at 
present ; and that the friction of the nervures produced a 
grating sound, as is now the case with the wing-covers of the 
females.^ A grating sound thus occasionally and accidentally 
made by the males, if it served them ever so little as a love-call 
to the females, might readily have been intensified throngh 
sexual selection, by variations in the roughness of the nervures 
having been continually preserved. 

In the last and third Family, namely the Acridiidte or 
grasshoppers, the stridulation, is produced in a very different 
manner, and according to Dr. Sciidder, is not so shrill as in the 
preceding Families. The inner surface of the femur (fig. 14, r) 
is furnished with a longitudinal row of minute, elegant, lancetr 
shaped, elastic teeth, from 85 to 93 in number ;" and these are 
scraped across the sharp, projecting nervures on the wing-covers, 
which are thus made to vibrate and resound. Harris" saya 
that when one of the males 
begins to play, bo first " bende 
" the shank of the hind-leg 
'■' beneath the thigh, where it 
" is lodged in a furrow de- 
" signed to receive it, and 
" then draws the leg briskly 
" up and down. He does not 
' play both fiddles t^ether, 
" but alternately, first upon 
" ono and then on the other." 
In many species, the base 
' of the abdomen ia hollowed 
out into a great cavity which 
is believed to act as a re- 
\ ii«niBed gounding hoard. In Pneu- 
mora (fig. 15), a S. African 
genus belonging to the same family, we meet with a new 

" Mr. Walsh also iaforms me " wing-covers ti^ther." 
that he has noticed that the famala *' Laniois, ibid. a. 113, 

of the Flatyphi/ilam ccncavum, " ' Insects ef Mew EDgiandj 

"whee captored ma^es a feehle 1842, p. 133. 
"gisUDg noise hf shuffling her 


Ceap, X 



and romtwkablo modification; ia tha males a small notched 
ridge projects obliquely from each side of the abdomen, 
against which the hind femora are rubbed.'* As the male is 
furnished with wings (the female being wingless), it is re- 
markable that the thighs are not rubbed in the usual manner 
against the wing-covers ; but this may perhaps be aceoimted for 
by the unusually smaO size of the hind-legs. I have not been 
ace of the thighs, which, judging 

flritleh Museum). Oppcr Bgure, nu 

from analogy, would be finely serrated. The species of Pneumora 
have been more profoundly modified for the Bake of stridulation 
than any other orthopterous insect; for in the male the whole 
body has been converted into a musical instrument, being 

"-Westwood,,' Modei-n Classification,' toI. 1. p. 463. 


288 The Descent of Man. Paut H. 

distended with, air, like a, great pellucid bladder, so as to 
Increase the reaonanoe. Mr. Trimen informs ma ttiat at the 
Cape of Good Hope these insects make a wonderful noiste 
during the night. 

In the three foregoing families, the females are almost always 
destitute of as efficient musical apparatus. But there are a few 
exceptions to this rule, for Dr. Gruber has shewn that both 
sexes ot Ephippiger vitium are thus provided; though the organs 
differ in the male and female to a cei-toiu eitent. Hence we 
cannot suppose that they have been transferred from the male 
to the female, as appears to havo been the case with the secondary 
sexual characters of many other animals. They must have been 
independently developed in the two eexes, which no doubt 
mutually call \o each other during the season of loTe. In most 
other Locustidie (but not according to Landois in Dectiens) the 
females have rudiments of the stridulatoty organs proper to the 
male; from whom it is probable that these have been transferred. 
Landois also found such rudiments on the under surface of the 
wing-covers of the female Achetidje, and on the femora of tho 
female Acridiidje. In the Hcanoptera, also, the females have the 
proper musical apparatus in a functionless state ; and we shall 
hereafter meet in other divisions of tho animal kingdom with 
many instances of structures proper to the male being present 
in a rudimentary condition in the female. 

Landois has observed another important fact, namely, that in 
the females of the Acridudae, the stridulating teeth on the 
femora remain throughout life in the same condition in which 
they first appear during the larval state in both sexes. In the 
males, on the other band, they !>ecome further developed, and 
acquire their perfect structure at the last moult, when the insect 
is mature and ready to breed. 

Prom the facts now given, we see that the means by which 
the males of the Ovthoptera produce their sounds are extremely 
diversified, and are altt^etber different from those employed by 
the Homoptera." But throughout the animal kingdom wo 
often find the same object gained liy the most diversified means ; 
this seems due to the whole organisation having undergone mul- 
tifarious changes in the course of ^es, and as part after part 
varied different variations were taken advantage of for tho 
eame general purpose. The diversity cf means for producing 
Bound in the three tionilies of the Orthoptera and in the 

" landois has recently found in niupteraj and Ihia is a smprlsiEg 

owiain Orthoptera rudimentary fact. Sie 'Zettschr. fUr wiasentth. 

raTietorea cloaelj similar to the Zoolog." B. iiii. Heft 3, 1871, j» 

Bo-.ra-prod ::cine urgans in the Ho- 348, 


Chap. S. Qrthoptera. 28q 

Honioptera, impres3<s the mind with the high importaiiee ol these 
BtructureB to the males, for the sake of calling or allitring the 
females. We need feel no surprise at the amoont of modification 
which tho Orthoptera have undergone in. this respect, as we now 
snow, from Dr. Scudder's remarkahle discovery," that there has 
Deen more than ample time. This naturalist has lately found 
a fossil insect in the Devonian formation of New Briinswiok, 
which is fumished with " the well-known tympanum or stridu- 
" latjng apparatus of the male Locustidaj," The insect, though 
in most respocts related to the Neuroptera, appears, as is so often 
the case with very ancient forms, to connect the two related 
Orders of the Neuroptera and Orthoptera. 

I haTe but little more to say on the Orthoptera. Some of the 
species are very pugnacious : when two male field-crickets 
{Qfyilua campestHs) are confined together, Oiey fight till one 
kills the other; and the species of Mantis are described as 
manceuTring with their sword-like front-limbs, like hussars with 
their sabres. The Chinese keep these insects in httle bamboo 
cages, and match them like game-cocks." With respect to 
colour, some exotic locusts are beautifully onamented; the 
posterior wings being marked with red, blue, and black; bnt as 
throughout the Order the seios rarely differ much in colour, it 
is not probable that they owe their bright tints to sexual 
selection. Ooospicnous colonrs may be of use to these insects, 
by giving notice that they are unpalatable. Thus it has been 
observed '" that a bright-coloured Indian locust was invariably 
rejected when offered to birds and Hzards. Some cases, however, 
are known of sexual differences in colour in this Order. The 
male of aa American cricket" is described as being as white as 
JTory, whilst the female varies from almost while to greenish- 
yeUow or dusky. Mr. Walsh informs me that the adult male of 
Spectrum /emoratum (one of the Phasmidje) " is of a shining 
"brownish-yellow colour; the adult female heii^ of a dull, 
" opaque, cinereous brown; the yonng of both seses being green." 
Lastly, I may mention that the male of one curious kind ot 
flrioket" is furnished with "a long membranous appendage, 
" which falls over tho lace like a veil ;" bat what its use may be, 
is not known. 

" 'TraD»acl.Ent.Soc.'3ra series, " The (Ecantlius nivalis. Karris, 

Tol. a, ("Journal of i'roecedings,' 'Insects of New England,' 1842, ji. 

p. 117.) 134. ThatwosBieaof ffi.pcflueW-^ 

« Wcstwood, 'Modern tlaaa. of of Europe differ, as I hear from 

Inwols,' toI. i, p. 427 ; for crickets, Victor Cams, in nearly the Mine 

p. 44.'!. manner. 

'■ Mr. Ch. Hnme, in ' Proc Enl. " PlatyUemniis : Weetwooi, 


290 The Descent of Man. PartjI. 

Order, NennrptJ'ra — Little need here be said, except as to 
colour. In the Ephemi-ndEe the seses often differ slightly in 
their obscure tints , *' but it is not probable that the males aie 
thus rendered attractive ti) the females. The Libellnlidffi, or 
dragon-flies, are ornamented with splendid green, blue, yellow, 
and Yonmliou metallic tints; and the seses often differ. Thus, 
as Prof. We'.twood remarks," the males of some of the 
Agrjonidffi, " are of a rich bine with black wings, whilst the 
" females are fine green with colourless wings." But in Agrion 
Mambuni these colours ate exactly reversed in tJie two seses." 
In the extensive N. American genus of Hetwrina, the males alone 
have a beautiful carmine spot at the base of each wing. In 
Anax Junius the basal part of the abdomen in the male is avivid 
ultramarine blue, and in the female grass-groon. In the allied 
genns Gomphus, on the other hand, and in some other genera, 
the sexes differ but little in colour. In closely-allied forms 
tlu'oughout the animal kingdom, similar cases of the seses 
differing greatly, or very little, or not at all, are of frequent 
occurrence. Although tiiere is so wide a difference in colour 
between the sexes of many Inbellnlida, it is often difficult to say 
■which is the more brilliant ; and the ordinary coloration of the 
two sexes is reversed, as we have jnst Been, in one species of 
Agrion. It is not probable that their colours in any case have 
been gained as a protection. Mr. MacLachlan, who has closely 
attended to this family, writes to me that dragon-flies— the 
tyrants of the insect-world — are the least liable of any insect to 
be attacked by birds or other enemies, and he believes that their 
bright colours serve as a sosnal attraction. Certain dragon-flies 
apparently are attracted by particular colours ; Mr. Patterson 
observed *" that the Agrionidje, of which the males are blue, 
settled in mmibers on the bine float of a fishing line ; whilst two 
other species were attracted by shining white colours. 

It is an interesting fact, first noticed by Sehelrer, that, in 
several genera belonging to two sub-families, the males on first 
emergence &om the pupal state, are coloured exactly Uke the 
females ; but that their bodies in a short time assume a con- 
spicuous milky-blue tint, owing to the exudation of a kind of oil, 
soluble in ether and alcohoL Mr. MacLachlan believes that in 
the v\sX& of Libdlula de^essa thischangeof colour does not occur 
until nearly a fortnight after the metamorphosis, when the sexes 
are ready to pair. 

" B. D. Walsh, the ' Pseudo-neu- indebted to thil nstnraliBt for ths 
rnptera of UliuMi,' ia 'Pros. Eat. following facts on Hetarina, An.ii, 
Soe. of Philadelphia,' 1862, p. Sei. dad Gomphus. 

»' Modern Class.' vol. ii. p. 37. "'Transact. Ent. Soc' vol. L 

" Walsh, ibi<l. i>. 381. I am IS36, p. Iiiii. 


OflAP. X. Hymenoptera. 291 

Certain species of Neurothetnis present, aocordii^ to Brauer, " 
A carious case of dimorphism, some of the females having ordinary 
wicRs, whilst others haTo them " very richly netted, as in the 
'■ males of the same speeies." Braner " explains the phenomenon 
■' on Darwinian principles by the supposition that the close 
" netting of the veins is a secondary sesnal character in the 
•' males, which has been abmptiy transferred to some of the 
" females, instead of, as generally occnis, to all of them." Mr. 
Ma«LachIan inibrms me of another instance of dimorphism 
in several species of Agrion, in which some individnals are of 
an orange colour, and tke^ are invariably females. This is 
probably a case of reversion ; for in the tnie libellnto, when 
the sexes differ in colonr, the females are orange or yellow ; 
so that supposmg Agrion to be descended from some primordial 
form which resembled the typical LibelluliB in its sexual cha- 
racters, it would not be surprising that a tendency to vary in 
this manner should occur in the females alone. 

Although many dragon-flies are large, powerful, and Berca 
inaects, the males have not been observed by Mr. MacLa«hlan to 
fight together, cxoepting, as he believes, in some of the smaller 
Bpecies of Agrion. In another group in this Order, namely, the 
Termites or white ants, both sexes at the time of swarming may 
be seen running aboat, " the male after the female, sometimes 
" two chasing one female, and contending with great eagemesa 
" who shall win the prize."'* The Atropos puhatorias is said 
to make a noiso with its jaws, which is answered by other 

Order, HymeaopUra. — That inimitable observer, M. Fabre,^" in 
describing the habits of Cerceris, a wasp-like insect, remarks that 
" fights frequently ensue between the males for the possession of 
" some particular female, who sits an apparently unconcerned 
" beholder of the stru^'le for supremacy, and when the victory 
" is decided, quietly flies away in company with the conqueror." 
Westwood " says that die males of one of the saw-flies (Tenthre- 
dinje) " have been found fighting together, with their mandibles 
' locked." As M. Fabre speaks of the males of Cerceris striving 
to obtain a particular female, it may be well to bear in mind 
that insects belonging to this Order have tlie power of rccognisiag 

" See abstract in the ' Zoological "See an interesting article 

Record ' for 1867, p. *50. ■ Uie Writinea of Fabre,' in ' Nai. 

" Kirby and Spence, ■ lotroducl. Uiat. Revien-,^ April 1862, p. 122. 
(<i E=toipology,' vol. ii. 181(1, jk 33. " 'Jounial of Protof KntMcolog 

'■' 2c!iMau, * Les Faculti^ Men- Sec.' Sept. 7th, 1863, p. IBS. 
tfllfc),' liC. Tum. i. p. 101. 


292 The Descent of Man. Paot 11 

each other after long IntetTals of time, and are deeply attached. 
For instance, Pierre Huber, whose accuracy no one doubts, 
separated some ants, and when, after an interral of foor months, 
they met others which had formerly belonged to the sanie 
conmiimity, they recognised and caressed one another with their 
antennjB. Had they been etrangers they would have fought 
Whether. Again, wben two communities engage in a, battle, the 
ants on the same side sometimes att!u;k each other in tht genei'al 
confusion, bnt they soon perceiye their mistake, and the one ant 
soothes the other.^ 

In this Order slight differences in colour, according to ses, are 
common, but conspicuous diffiirocoeE are rare except ir the 
family of Bees; yet both scses of certaingroupsare so brilliantly 
coloured— for instance in Chrysis, in which Termdlion and 
metallic greens prevail— tliat we are tempted to attribute the 
result to sesua! selection. In the Ichneumonidffl, according to 
Mr. Walsh,™ the males are ahnost uniTersaUy lighter-coloured 
than the females. On the other hand, in the Tenthredinidte the 
males are generally darker than the females. In the Siricidse 
the seses frequently differ; thus the male of SirBe j'lwemcus is 
banded with orange, whilst the female is dark purple ; but it ia 
difQcnlt to say which sex is the more ornamented. In Trernex 
rdliimh(s the female is much brighter-coloured than the male. 
I am informed by Mr. P. Smith, that the male ants of several 
species are black, the females being testaceous. 

In the femily of Bees, especially in the solitary species, as I 
hear from the same entomologist, the seses often differ in colour. 
The males are generally the brighter, and in Bombns as well as in 
Apathus, much more variable in colour than the females. In 
Antliophnra rettisa the male is of a rich fulvoi\8-brown, whilst 
the female is quite black : so are the females of several species 
of Sylocopa, the males being bright yellow. On the other hand 
the females of some species, as of AndrccTia. fulva, are much 
brighter-coloured than the males. Such differences in colour 
can baldly bo accounted for by the males being defenceless and 
thna requiring protection, whilst the females are well defended 
tiy their stings. H, Miiller, *• who has particularly attended to 
the habits of bees, attributes these differences in colour in chief 
part to semal selection. That bees have a keen perception of 
colour is certain. He says that the males search eagerly and 
fight for the possession of tho females ; and he accounts through 

^ P. Huber, ' Recherches 6nr lei Philadelphia,' 1SS6, pp. "ftfunsfl. 
MatH d« Fourmis," 1810, pp. 150, *■ ' Anwenduag der llsrsinstten 

166. Lehre anf Biinwa.' Van. a. o. 

^ ' Pwe. Entomolog. Soc. of JaLi^. iiii. 


CtTiP. X. Hymenoptera. 293 

ench contests for the mandibles of the males being in certain 
speciea larger than those of the femaies. In some caees the 
males are far more numerous than the females, either early 
in the season, or at all times and places, or locally ; whereas tlie 
females in other cases are appai'ently in oxcees. In some species 
Uie more beantifol males appear to have been selected by the 
females ; and in others the more beautiful females by the males. 
Consequently in certain genera (Mliller, p. 42), the males of the 
several species differ much in appearance, whilst the female ate 
almost indistinguishable; in other genera the reverse occurs. 
H. Miiller belieTee (p. 82) that the coloura gained by one ses 
through sexual selection have often been transferred in a variable 
degree to the other sex, just as the pollen-eoUeoting apparatus 
of the female ha^ often been translbrred to the male, to whom 
it is absolutely useless." 

MvMUa Ewopsea makes a stridulatiog noise ; and accoriiing to 
Gonreau " both sexes have this power. He attributes the sound 
to the friction of the third and preceding abdominal segments, 
and I find that these surfaces are marked with very fine con- 
centric ridges ; but so is the projecting thoracic collar, into which ■ 
the head articulates, and this collar, when scratched with the 
point of a needle, emits the prox>er sound. It is rather surprising 
that both sexes should have the power of stridulating, as the 
male is winged and the female wingless. It is notorious that 
Bees express certain emotions, as of anger, by the tone of their 
humming; and accordii^ to H. Miiller (p. 80), the males of 
e species make a peculiar singing noise whilst pursning the 

•' M.Perrier in "nis article' la S^ male grandfathers? To take a cise 

lectionstinelled'apreaDarwin'CRe- witt oviiiiary animals as nearly 

TueScienCif]<[u«,'Feb. 18T3, p.SSSX parallel v» possible: if a female ot 

without apparently having refleoteil any white quadruped or bird were 

ranch on the subject, objects that as cTossed hy a male of a blacli breed, 

the males of sodol beef are kuowa and the male and female ofispiin' 

Vt be produced JVom unfertilised vere paired together, will it be 

oTa, they could not transmit new pretended that the grandchildren 

chsracters to their male offspring, would not inherit a tendency to 

This is an eitraordinarj objection, blackness from their male graud- 

A female bee fertilised by a male, father P The acquirement of new 

which presented some character fii- characters bj the sterile worker-bees 

dlltating Uie nnion of the seies, or is a much more ditlicult case, bnt I 

rendering him more attractive to have endeavoured to show in my 

the female, would lay eggs which 'Origin of Species,'liow thesBsterilj 

wonld produce only females; but beings are subjected to thepower ol 

these young females wonld neit natural selection, 

year produce males, and will ft be •' Quoted by Wert wood, 'Modern 

pretended that such males wouM Class, of Insects,' vol. ii. p. 31* 
DO* inherit the characters of their 


294 ^•^'^ Descent of Man. Past II 

Order, Coleoptera (Beetles).— Many beetles are coloured so &e 
to resemble the surfaces whicL they habitually frequent, ami 
they tlms escape detection by their enemies. Otter species, for 
instance diamond-beetles, ore ornaoiented with Bplendid colours, 
which are often arranged in stripes, spots, crosses, and othef 
elegant patterns. Such colours can hardly serve directly as a 
protection, escept in the case of certain flower-feeding species ; 
bnt they may serve as a waminK or taeans of reci^nitioD. on the 
same principle as thn phosphorescence of the glow-worm. As 
with beetles the colours of the two sexes are generally alike, we 
have no evidence that they have been gained through sexual 
seleotion; but this is at least possible, for thej may have been 
developed in one sex and then transferred to the other ; and 
this view is even in some degree probable in those groups which 
possess other well-marked secondary sexual characters. Blind 
beetles, which cannot of course behold each other's beauty, 
never, as I hear from Mr. Waterhouse, jun., exhibit bright 
colours, though they often have polished coats; but the expla- 
nation of their obscurity may be that they generally inhabit 
caves and other obscure stations. 

Some Longicoms, especially certain Prionidie, offer an excep- 
tion to the rule that the sexes of beetles do not differ in colour. 
Most of these insects are lai^e and splendidly coloured. The 
males in the genus Pyrodes," which I saw in Mr. Bates's col- 
leetioii, are generally redder but rather duller than the females, 
the latter being coloured of a more or less splendid golden-green. 
On the other hand, in one speciea the male is golden-green, the 
female being richly tinted with red and purple. In the genus 
Esmeralda the sexes differ so greatly in colour that they have 
been ranked as distinct species ; in one species both are of a 
beautiful shining green, but the male ha^ a red thorax. On the 
whole, as far as I could jndge, the females of those Prionidffl, in 

" Pyrodea jxilcherrmms, in 

the family of Longicoma. M 

which th«Beiea differ coUEpicuonsly. 

R. Trlmen and Waterhouse, 

nr« been deecribed by Mr. Bates in 

inform me of t"0 Lnmelli 

■Transact. Ent. Soo." 1869, p. 50. 

vli., a PeritriehU and Tiichiu 

I will apewfy the few other eases in 

male of the latter being 

which 1 bave heud of s difTerence 

obscurely coloured than the fe 

in colons between thfl Mxea of 

In TVJ/u! elongatas the male is 

t.eetl«L Kirby and Spenee ('In- 

and the female alwaje, as 

Irodoct. to Entomology,' vol. iii. p. 

Wieved, of a dark blue coloor 

301) mention a Cantharis, Metoe, 

a red thorai. The male, sL 

HhBgium,sad the ZtpCura teaiacea; 

OraodaCHa atra, as I hear fro 

the male of the latWr beinii tes- 

Walsh, is black, the feinale 

taceons, with a black thori., and 

so-called 0. ruJicoWri) havi 

the female of a dull red all ovei. 

■ufous thorat. 

These two Utter beetles beloag to 


Cbap. X. 



■whdcli the sexes differ, ate coloured more richly than the males, 
and this does not accord with the common rule in regard to 
colour, when acquired through sexual selection. 

A most remarkable distinction between the sexes of many 
beetles is presented by the great boms which rise from the head, 
thorax, and clypeus of the males; and in some few cases from 
the under surface of the body. These horns, in the great family 
of the Lamellicoms, resemble those of ■various quadrupeds, such 
as stags, rhinoceroses, &c., and are wonderful both from their 
eize and diversified shapes. Instead of describing them, I have 
given figures of the males and females of some of the more re- 
markable forms. (Figs. 16 to 20.) The females generally ex- 
hibit rudiments of the horns in the form of small knobs or 
ridges ; but some are destitute of even the slightest rudiment. 
On the other hand, the horns are nearly as well developed in the 
female as in the male of Fhwiueas lancifer; and only a little less 
well developed in the females of some other species of this genus 
and of Copris, I am informed by Mr. Bates that the horns do 
not differ in any manner corresponding with the more important 
characteristic differences between the several subdivisions of tho 
family : tlios within the same section of the genus Onthopbague, 
there are species which have a single horn, and others which 
have two. 

fcns le. Chiflcosolnii 


296 The Descent of Man. Paet 11. 

F%. \1. Copria teldis. (Left-lmnil flgnre^ rat 

Jr^^ Ji^. 

Chap. X CoUoptera. %tyj 

In almost all cases, the horns are remarkable from their ex- 
cessive variability; so that a graduated series can be formed, 
from the most highly deTeloped males to otiiers eo d^enerale 
that they can barely be distingtiished from tlie females. Mr. 
Walsh" found that ia Phrmceus caravan, the boms were thrice as 
long in some males as in others. Mr. Eaftes, after exomijiu^ 
above a hundred males of Onthophagvt rangi/er (Bg. 20), thought 
that he had at last discovered a species in which the horns did 
not vary ; but fnrther research proved the contrary. 

The extraordinary size of the boms, and their widely different 
Btrnctiire iu dosely-aUied forms, indicate that thej have been 
formed for some pnrpose ; but their excessive variability ia the 
males of the same species leads to the inference that this purpose 
cannot be of a definite nature. The horns do not show marks of 
friction, as if used for any ordinary work. Some authors sup- 
pose'" that as the males wander about much more than the 
females, they require horns as a defence against their enemies ; 
but HS the horns are often blunt, they do not seem well adapted 
for defence. The most obvious conjectnre is that they are ased 
by the males for fightii^ together; but the mfdes have never 
been observed to fight; nor could Mr. Bates, after a careful 
examination of numerous speciee, find any sufBcient evidence, ia 
their mutilated or broken condition, of their having been thus 
used. If the males had been habitual fighters, the size of their 
bodies would probably have been increased through sesnal 
selection, so as to have exceeded that of the females; bat 
Mr. Bates, after comparing the two sexes in above a hundred 
species of the Copridie, did not find any marked difTerence in 
this respect amongst well-developed individuals. In Lethrus, 
moreover, a beetle belonging to the same great division of the 
Lamellicorns, the males are known to fight, but are not provided 
with horns, though thfeir mandibles are much lai^er than those 
of the female. 

The conclusion that the horns have been acquired as ornaments 
is that which bast agrees with the fact of their having been so 
immensely, yet not fixedly, developed, — as shewn by their extreme 
variabiKty in the same species, and by their extreme diversity in 
closely-allied species. This view will at first appear extremely 
improbable; but we shall hereafter find with many animals 
standi!^ much higher in the scale, namely fishes, amphibians, 
reptiles and lards, that various kinds of crests, knobs, horns and 
(lombs have been developed apparently for this sole purpose. 

The males of Onilia furci/^ (fig. 21), and of some ottior 

" ' Proo. Entomolog. So«. of *' Kirby and Spenoe, 'iDtrodnct. 
PUIadclphJa, 1S61, p. ass Eotomolog.' vol. IIL p. 300, 



s Descent oj Man. 


species of the genus, are fnrnished with singular projections on 
their anterior femora, and with a great fort or pair of homs on 
the lower surface of the thoras. jTidging 
from other insects, these may aid the male 
in clinging to the female. Althoegh the 
males have not even a trace of a hora on 
the upper surface of the hody, yet the fe- 
males plainly exhibit a rudiment of a single 
horn on the head (fig. 22, a), and of a crest 
(b) on the thoras. That the slight thoracio 
f tt « crest in the female is a rudiment of a pro- 

T" r » jeetion proper to the male, though entirely 

Fig 21. Oqitia turcifer, absent ]n the male of this particular species, 
n'eaiV''''* ** '""" ^ ^^ cleaT for the female of B^tbas hiatm fa 
frenus which comes nest to Onitis) has a 
Eimilai slight crest on the thoras, and the male bears a great 
prtgection in the same sitnation. So, again, there can hardly bo 
a doubt that the bttle point (a) on the head of the female Onitis 

Icft-hsnd figure, 

farci/er, as well as on the head of the females of two or three 
allied species, is a rudimentary representative of tie cephalic 
horn, which is common to the males of so many LameUicom 
beetles, as in PhaEEeus (fig. 18). 

The old behef that rudiments have been created to complete 
the scheme of nature is here so far from holding good, that we 
have a complete inversion of the ordinary state of things in the 
family. We may reasonably suspect that the males originally 
bore homs and transferred them tc the females in a rudimentary 
condition, as in so many other Lamellicoms. Why the males 
subsequently lost their horns, we know not; but this may have 
been caused through the principle of compensation, owing to 
the development of the lai^e horns and projections on the lower 
surface ; and as these are confined to the males, the rudiments 
of the upper homs on the females would not have been 





The cases hitherto given refer to the Lamellicoms, bat the 
males of some few other heetles, belonging to two widely distinc'i; 
groups, namely, the CurculionidsB aad Ktaphylinidse, are fur- 
nished with horns— in the former on the lower surfiice of the 
body," in tlie latter on the npper surfece of the head and thorax. 
In the StaphylinidEB, the horns of the males are extraordinarily 
variable in the same species, just as we have seen with the 
Lamellicorns. la Siagonium we have a case of dimorphism, 
for the males can be divided info two seie, differing greatly 
in the size of their bodies and in the development of their 
horns, without intermediate gradations, in a species of Bledius 
(fig. 23), also belonging to the Staphylinidse, Professor Westwood 

states that, " male specimens can be found in the same locality 
" in which the central horn of the thorax is very lai^e, but the 
" horns of the hea^ quite rudimental ; and others, in which the 
"thoracic horn is much shorter, whilst the protuberances on 
" the head are long."" Here we apparently have a case (rf 
compensation, which throws fight on that just given of the 
Buppoeed loss of the upper homa by the males of Onilu. 

Law of Battle.— Bam& male beetles, which seem ill-fitted for 
fighting, nevertheless engage in conflicts for the possession of 
the females. Mr, Wallace" saw two males of Leplorhynchus 
aTigtislaf-us, a linear beetle with a much elongated rostrum, 
" fighting for a female, who stood close by busy at her borinp, 
" They pushed at each other with their rostra, aad clawed and 
" thumped, apparentlyin the greatest rage." The smaller male, 
however, " soon ran away, acknowledging himself vanquished." 
In some few cases male beetles are well adapted for fighting, by 
possessing great toothed mandibles, much larger than those of 
the females. This is the case witti the common etag-beetio 
(i««Mi(w cereus), the males of which emerge from the pupal 
Btafo about a week before the other sex, so that several may 
) female. At this sesBon thoy 

often be seen pursuing the s 

" Kirby and Spence, ' Introduct, 
Entomolog.' vol. iii. p. 329. 

" ' Modern ClaBBification of In- 
sects,' vol. i. p. 172 ! Siagoninin, 
p. 173. In the British Museum I 
noticed one ni!i1e epecimen of Sia- 

' The Malay Archipelago,* vol. 
J89,p.276. Eiley, Sisth ' Eaport 
(sects of Missoori," 1874, p. iii 


300 Tlie Descent of Man. Past H 

engage in fierce conflicts. "When Mi. A. H, Davis" enclosed 
two malee with one female in a bos, the larger male severely 
pinched the smallor one, until he resigned his pretensions. A 
friend informs me that when a hoy he often put the males 
together to see them flght, and he noticed that fhey were much 
bolder and fiercer than the females, aa with the higher animals. 
The males would seize hold of his finger, if held in front ot 
them, but not so the females, although they have stronger 
]aws. The males of many of the Lucanidie, as well as of the 
above-mentioned Leptorhjnchus, are lai^r and more powerful 
insects than the females. Tbe two sexes of Lethrvs ceplialotia 
(one of the LarecllioomB) inhabit the same burrow; and the 
male has lai^r mandibles than the female. If, during the 
breeding-season, a strange male attempts to enter tbe burrow, 
he is attacked; the female does not remain passive, but ctoees 
the moutJi of the burrow, and encourages her mate by con- 
tinually pushing bim on from behind ; and the battle lasts until 
the a^ressor is killed or runs away." The two sexes of another 
Lamellicorn beetle, tte AleuAvs cjcnin'cosus, hve in pairs, and 
Beem much attached to each other ; the male escitcs the female 
to roll the balls of dung in which the ova are deposii«d ; and if 
she is removed, he becomes much agitated. If the male is 
removed the female ceases ail work, and asM.Briilerie'" believes, 
would remain on the same spot until she died. 

The great mandibles of the male LucanidtB are extremely 
variable both in size and Btructure, and in this respect resemble 
the horns on the head and thorax of many male Lamellicoms 
and Staphjlinidfe. A perfect series can be formed from the 
best-provided to the worst-provided ot degenerate males. Al- 
though the mandibles of tbe common stag-beetle, and probably 
of many other species, are used as efficient weapons for fighting, 
it is doubtful whether their great size caa thus be accounted 
for. We have seen that they are used by the Liuxirms elaphta 
of N. America for seizing the female. As they are so con- 
spicuous and so elegantly branched, and as owing to their great 
length they are not well adapted for pinching, the suspicion 
has crossed my mind that they may in addition serve as an 
.-imament, like the horns on the bead and thorax of the various 
species above described. The male Chianognathus Grantii of 
8. Chile — a splendid beetle belonging to the same family— baa 

•• 'Entomological Mag.izine,' voL " Quoted from Fischer, In ' Dirt, 

1. 1833, p. 82. See also on the Class. d'Uiat. Nat." torn. i. p, 324. 
conflicts of tbia Bpec!e3, Kirby and " ' Aaa. Soc. Entomolog, France, 

Spenee, iMiL vol. iii, p. 314; ani 186G, as quoted In 'Jonrnal of 

Wfstwwrf, IMd, vol, i, p. 187. Trdvel,' ly A. Mnrray, 1888, p. 1S5. 


Chap. X. 


enormously developed mandibles (fig. 24) ; he is bold and pug- 
nacious; when threatened he faces round, opens his great jaws, 
and at the same time stridulates loudly. 
But the mandibles were not strong 
enoi^h to pinch my finger so aa to 
cause actual pain. 

Sexual selection, which implies the 
possession of considerable perceptive 
powers and of strong passions, seems 
to have been more effective with the 
Lamellicoms than with any otbec 
family of beetles. With some species 
the males are provided with weapons 
for ^hting; some hve in pairs and 
show mutual affection; many have 
the power of strldulatii^ when excited ; 
many are furnished with the most ex- 
traordinary horns, apparently for the 
sake of ornament; and some, which 
are diurnal in their habits, are gor- 
geonsly coloured. Lastly, several of 
the largest beetles in the world belong 
to this family, which was placed by 
Linnieus and Fabricios at the head of 
the Order." 

Stridulatinff organs. — Beetles belong- 
ii^ to many and widely distinct 
famiUes possess these organs. The 
sound thus produced can sometimes 
be heard at the distance of several feet 
or even yards,'" but it is not comparable 
with that made by the Orthoptera. 
The rasp generally consists of a narrow, 
slightly-raised surface, crossed by very 
fine, parallel ribs, sometimes so fine as 
to cause iridescent colours, and having 
a very elegant appearance under the 
microscope. In some cases, as with 
Typhceus, minute, bristly or scale-like ' 
prominences, with which the whole 

surrounding surface is covered i_ _ j, 

lines, could be traced passing into the ribs of the ra.sp! The 

" Westwood, ' Moiei'Q Class.' Cureulioi 
rol. i p. 184, Mat. Hist 

" WollastoD, ' On certain Musical 

riF.S^. CliEaeognatbusgrrm^i, 
rednwd. Upper flRnre, male; 
lower Bgure, female. 

. approximately parallel 


e Descent of Man. 


tiansitJon takes place by their becomms confluent and ritiight, 
and at the same time more prommtait and smooth A hanj 
ridge oa an adjoining part ot the bsdy serves as the scrapcf 
for the rasp, but this scraper m some ca^es has been specially 
modified for the purpose. It is rapiiily movei' across the rasp, 
or conversely the rasp across the sciap i 

the carrion-bettles (Necrophorus) two parallel rasps (r, fig. 25) 
stand on the dorsal surface of the fifth abdominal segment, eacli 
rasp" conaistii^ of 126 to 140 fine ribs. These ribs are scraped 
against the posterior mai^ns of the elytra, a small portion of 
■which projects beyond the general outline. In many Crioceridie, 
and in Clythra 4r^ndata (one of the Chrysomehdae), and m some 
Tenebrjonidse, &c.," the rasp is seated on the dorsa] apex of the 
abdomen, on the pygidium or pro-podium, and is scraped in 
the same manner by the elytra. In Hetcrocerus, which belongs 
to another family, the rasps are placed on tbe sides of the 
first abdominal segment, and are scraped by ridges on the 
femora." In certain Curculionidte and Carabidas," tlie parts 

" Lsndois, 'Zcitsohrift fur wiss. 
Zoolog.' B. :[vii. 1867, s. 137. 

" I am greatly indebted to Mr. 
G. R. Crotch for having sent me 
maoy prepared specimens of rarions 
beetles belonging to these thiea 
femilies and to oiJier3,aswell as for 
ralnable information. He believes 
that the power of stridnlation in 
the Ciythra has not been previously 
observed. I am also much 'nd bted 
to Mr. E. W. Janson, fo nfom a 

that my son, Mr. F. Darw n finds 
that Dermestes murimis stndulat 
hut he searched in vai lo the 
appiTBtos. Scolytus has lat ly 

been described by Dr. Chapman as 
a stridulator, in the 'Entomolo- 
gist's Monthly Magazine,' vol. vi. p. 

" Schiodte, translated in ' Annals 
and Mag. of Hat. Hist.' vol. ir. 
1867, p. 37. 

" Westring has described (Kroyer, 
' Natorhist. Tidstrift,' B. ii. 1848- 
49, p. 334) the stridulating organs 

n these two, as well as in c^er 
fara 1 es. In the Carabidx I have 

lam ned Elaphrus viiginosus and 
Bleth sa mtdlipuaclata, sent to me 
by M . Crotch. In Bletliisa the 
trans erse ridges on the furrowed 
bo der of the ^idomina] segment da 


Ghaf. X. 



are completely reversed in position, for the rasps are seated on 
tiie inferior surface of the elytra, near their apices, or atong 
their outer margics, and the edges of the abdomiital segmenfe 
serve as the scrapers. In PehMus Hermanni (one of Dytiscidw 
or water-beetles) a strong ridge runs parallel and near to the 
sutural margin of the elytra, and is crossed by ribs, coarse tn 
the middle part, but becoming gradually finer at both ends, 
especially at the upper end ; when this insect is held under 
water or in the air, a stridulatii^ noise is produced by the 
extreme horny margin of the abdomen being scraped against 
the rasps, la a groat number of loag-borned beetles (Longi- 
comia) the organs are situated ijuite otherwise, the rasp being 
on the meso-thorax, which is rubbed against the pro-tborax; 
Landois counted 238 yery fine ribs on the rasp of Cerambyx 

Many Lamellicoms have the power of stridulati'ug, and the 
organs differ greatly in position. Some species etridulate Tery 
loudly, so that when Mr. F, Smith caught a Trox sabulosus, a 
gamekeeper, who stood by, thought he had caught a mouse ; 
but I failed to discover the proper oi^ns in 
this beetle. In Geotmpes and Typhraus a 
narrow ridge runs obliquely across (r, fig. 26) 
the coxa of each hind-leg (having in 0. ster- 
corarius 8i ribs), which is scraped by a 
specially projecting part of one of the ab- 
dominal segments. In the nearly allied Copria 
lunaris, exy excessively narrow fine rasp runs 
along the sntural margin of the elytra, with 
another short rasp near the basal outer mar- 
gin; but in some other Coprini the rasp is 
seated, according to Leconte,'* on the dorsal 
Burfece of the abdomen. In Oryctes it is 
seated on the pro-p^dium; and, according to 
the same entomologist, in some other Dynastini, ^ ^ 
on the under surface of the elytra. Lastly, 
Westiiag states that in Omaloplia brunnea the 
rasp is placed on the pro-sternum, and the " FmuV. "' (7'Tiwi 
scraper on the meta-stemum, the parte thus ""' '^'"^' 
occupying the under surface of the body, instead of the upper 
surface as in the Longicoms. 

We thus see that in the different coleopterous femiliea the 

not, as &r as I could judge, come of Illinois, for having sent me si. 

into plav in scraping tiie rasps on tracts from Leconle's ' IntroductioE 

the elytra. tu Eotomology,' pp. 101, 143. 
■ •' 1 am indebted to Mr. Walsh, 


304 T}ie Descent of Man. Pai:t n 

Etridulating organs are wonderfully diversified in position, but 
not mucb in structute. Withiu the same family some species 
are profided with theso organs, and others are destitute of them. 
This diTcrsity is inteihgible, if we suppose that originally vafioua 
beetles made a shuffling or hissing noise by the rubbing together 
of any hard and rough parts of their bodies, which happened tj> 
be in contact i and that iiom the noise thus produced being in 
some way useful, the rough surfaces were gradually developed 
into I'egulai stridulating oi^ans. Some beetles as they raoT«, 
now produce, either intentionally or unintentionally, a shuffling 
n lise, without possessing any proper organs for the purpose, 
Mr, Wallace informs me that the Huckirus lonijimanus (a 
Lamellicorn, with the anterior legs wonderfully elongated in the 
male) " makes, whilst moTiug, a low hissing sound by the pro- 
" trusion and contraction of the abdomen; and when seized it 
" produces a grating sound by rubbing its hind-legs against the 
" edges of the elytra." The hissing sound is clearly due to a 
narrow rasp running along the sutural margin of each elytron ; 
and I could likewise make the grating sound by rubbing the 
shagreened surface of the femnr against the granulated margin 
of Uie corresponding elytron ; but I could not here detect any 
proper rasp ; nor is it likely that I could have overlooked it in 
so large an insect. After examining Cycbrus, and reading what 
Westring has written about this beetle, it seems very doubtful 
whether it possesses any true rasp, though it has the power of 
emitting a sound. 

From the analogy of the Orthoptera and Homoptera, I 
expected to find the stridiilating organs in the Coleoptera 
differing according to sei; but Landois, who has carefully 
examined several species, observed no such difference ; nor did 
Westring ; nor did Mr. G. E. Crotch in prepajing the many 
specimens which he had the kindness to send me. Any difference 
in these oi^ns, if slight, would, however, be difficult to detect, 
on account of theii great variability. Thus, in the first pair of 
specimens of Necrophottts humator and of Peltiias which I ex- 
amined, the rasp was considerably larger in the male than in 
the female; but not so with succeeding specimens. In Qe/i- 
triipfs $teTcorarija the rasp appeared to me thicker, opaquer, 
and more prominent in three males than in the some number of 
females; in order, therefore, to discover whether the sexes 
differed in their power of stridulating, my son, Mr. F. Darwin, 
collected fifty-seven living speeimens, which he separated info two 
lots, according as they made a greater or lesser noise, when held 
m the same manner. He then examined all these specimens. 
Bud found that the males were very nearly in the same pj'oportion 


Chap. S, CoUoptera. 305 

to the females in both the lots. Mr. F. Smith haa feept alive 
iiniiierous specimens of Moiwjfnckas pseudacvri (Gurculionidie), 
»uA is couTinced that both seies etridulate, and apparently in 
on equal degree, 

KeTertteteB, the power of stridtdating is certainly a sosuat 
character in some few Coleoptera. Mr. Crotch discovered that 
the males alone of two sjieeies of Heliopathes (TcnebrionidM) 
possess stridulating organs. I examined five males of H. gihlmn, 
and in all these there was a well-developed rasp, partiaDy 
divided into two, on the dorsal surface of the terminal abdominal 
fif^nent ; whilst in the same number of females there was not 
even a rudiment of the rasp, the membrane of this segment 
beii^ transparent, and much tliinner than in the male. In 
B. cribmtostriatua the male has a similar rasp, excepting that it 
is not partially divided into two portions, and the female is 
completely destitute of ttis oi^n ; the male in addition has on 
the apical margins of the elytra, on each side of the suture, 
three or four short longitudinal ridges, which are crossed by 
extremely fine ribs, parallel to and resembling those on the 
abdominal rasp ; whether these ridges serve as an independent 
rasp, or as a scraper for the abdominal rasp, I could not decide : 
the female exhibite no trace of this latter stmcture. 

Again, in ibree species of the LameUJcom genus Oryotes, we 
have a nearly parallel case. In the females of 0. gryph-us and 
nasioomia the ribs on the rasp of the pro-pygidinm are less 
continuous and less distinct than in the males ; but the chief 
difference is that the whole upper surface of this segment, when 
held in the proper light, is seen to be clothed with hairs, which 
are absent or are represented by excessively fine down in the 
Kuiles. It should be noticed that in all Coleoptera the efiective 
part of the rasp is destitute of hairs. In 0. eenegaleitm the 
difference between the sexes is more strongly marked, and this 
is best seen when the proper abdominal segment is cleaned and 
viewed as a transparent object. In the female the whole surface 
is covered with little separate crests, bearing spines ; whilst in 
the male these crests in proceeding towards the apex, become 
more and more confluent, regular, and naked; so that three- 
fourths of the segment is covered with extremely flne parallel 
ribs, which are quite absent in the female. In the females, 
however, of all tiiree species of Oryctes, a slight grating or 
itridulating sound is produced, when the abdomen of a softened 
specimen is pushed backwards and forwards. 
In the case of the Heliopathes and Oryctes there can hardly 
, be a doubt that the males stridnlate in order to call or ta 
excite the females; but with most beetles the stridnlation 


3o6 The Descent of Man. PinT IL 

ftpparently serves both sexee as a nmtual call. Beetles stridu- 
late Tmder variona emotions, in the same maTiiier as birds Ms» 
their voices for many purposes besides singing to their mates. 
The great Chiasognathus stridulates in anger or defiance; many 
species do the same from distress or fear, if held so that they 
cannot escape; by striking the hollow stems of trees in the 
Canary Islands, Messrs. Wollaaton and Croteh were able to 
discover the presence of beetles belonging to the genus AcalleB 
by their stridulation. Lastly, the male Alcuchus stridnlatea to 
encourage the female in her work, and from distress when she 
is removed." Some naturalists beheve that beetles make t h i i 
noise to frighten away their enemies ; but I cannot think that 
a quadruped or bird, able to devour a large beetle, would 
be frightened by so slight a sound. The belief that the stridu- 
lation serves as a sexual call is supported by the fact that death- 
ticks {AtiMMm, tessellatum) are well known to answer each 
other's ticking, and, as I have myself observed, a tapping noise 
artificially made. Mr. Doubleday also informs me that he 
has sometimes observed, a female ticking," and in an hour or 
two afterwards has found her united with a male, and on one 
occasion surrounded by several males. Finally, it is probable 
that the two sexes of many kinds of beetles were at first 
enabled to find each other by the slight shuffling noise produced 
by the rubbing tc^ether of the adjoining hard parts of their 
bodies; and that as those males or females which made the 
greatest noise succeeded best in finding partners, rugosities on 
various parts of their bodies were gradually developed by means 
of sexual selection into true atridulating organs. 

" M, P. d« U BruleriB, ai quoted 


' Zeitechrirt 

far wissen. 

In 'Journal of Travel,' A. Murray, 


B. ,Vii. B. 1 

31. Oliver 

fol. i. 1868. p. 13.S. 

Kirb; and 

Introduet.' vol 

. ii. p. 395) 

" the noisB is produced by the in- 

that the 

female of Pii 


" >ect r«isine itself od Iu legs as 


d Mnnd by 

" high as it can. and then striking 


her abdnmen 

against any 

"its thorai five or sii timas, in 

hu-d Eub 

stance." and th 

^t the m.-ili. 

" rapid Buccession, against the snb- 

It to this call, . 

wja atMn«U 


Butterflies and Moths. 

Ikbbctb, contiimed. — Oedbk Lepidopteka, 


Courtship of bntterilies— Battles— Ticking noise— Colours common to 
both Mies, or more biillmnt in the mnics— £iampl«s — Not doe to the 
direct action of the conditions of life — Coionrs adapted for protection — 
Colonrs of moths— Display— Perceptiie powers of the Lepidoptera — 
Variability — Causes of the difference in colonr between the males and 
females — Mimicry, female bottarfiiea more brilliantly colonred than 
the males — Bright colours of oaterpiliars — Summary and concluding 
remarks on the secondary sexual characters of insects — Birds and insects 

In this great Order the most intereatiiig points for us are the 
diflferencea in colour between the seses of the same species, and 
between the distinct epecies of the same genus. Nearly the 
whole of the following chapter wiU be devoted to this subject; 
but I will first make a few remarks on one or two other points. 
Severftl males may often be seen pursuing and crowding round 
the same female. Their courtship appears to be a prolonged 
aSaii, for I have frequently watched one or more males pirouet- 
ting round a female until I was tired, without seeing the 
end of the courtship. Mr. A. G. Butler also informs me that 
he has several times watched a male courting a female for a full 
quarter of an hour; but she pertinaciously refused him, and at 
last settled on the ground and closed her wings, so as to escape 
from his addresses. 

Although butterflies are weak and fragile creatures, they are 
pugnacious, and an Emperor butterfly' has been captured with 
the tips of its wings broken from a conflict with another male. 
Mr. Collit^wood, in speaking of the frequent battles between the 
butterflies of Borneo, says, " They whirl round each other with 
" the greatest rapidity, and appear to be incited by the greatest 
" ferocity." 

The Ageronia feroriia makes a noise like that produced by a 
toothed wheel passing under a spring catch, and which can be 
heard at the distance of several yards : I noticed this sound at 
Eio de Janeiro, only when two of these butterflies were chasing 
each other in an irregular course, so that it is probably made 
during the courtship of the seses." 

' Apaiara Xrit: ' The Entomolo- Naturalist,' 1868, p. 183. 
gtst's Weekly Intelligence,' 18S9, p. " See my ' Journal of Eesearches, 

13S. For the Borusan Bntterflies, 1845. p. 33. Mr. Doubleday ha> 

fee C. Collicgwoad, ' Bambles of a detected (' Proc Ent. Soc' March 



The Descent of Jl 

Some moths also produce sounds ; for instance, the males of 
Tlieaiphora foocu. On two occasions Mr. P, Bnohanaa Wiita' 
beard a sbaip quick noise made bj the male of HyhpMla 
prasinana, and which he believes to be produced, as in Cicada, 
hj an elastic membrane, furnished with a. muscle. He quotes, 
also, Guenee, that Setina prodnces a sotmd like the ticking 
of a watch, apparently by the aid of " two largo tympanifonn 
" vesicles, sitiwited in the pectoral region ; " and these " are much 
" more developed in the male than in tie female." Hence tie 
sound-producing organs in tba Lepidoptera appear to stand in 
some relation with the sexual functions. I have not alluded 
to tbe well-known noise made by the Death's Head Sphinx, for 
it is generally heard soon after the moth has emerged from 

Girard has always obHerved that the musky odour, which is 
emitted by two species of Sphinx moths, is peculiar to the males ;' 
and in the higher classes we shall meet with many instances of 
tbe males alone beii^ odoriferons. 

Every one must have admired the extreme beauty of many 
butterflies and of some moths ; and it may be asked, are their 
colours and diversified patterns the result of the direct action of 
the physical conditions to which these insects have been exposed, 
without any benefit boiag thus derived ? Or have successive 
variations been accumulated and determined as a protection, 
or for soma unknown purpose, or that ono sex may be at- 
tractive to the other ? And, again, what is the meaning of the 
colours being widely different in the males and females of 
certain species, and alike in the two sexes of other species of the 
same gennB? Before attempting to answer these questions a 
body of facts must be given. 

With our beautiful EngUsh butterflies, the admiral, peacock, 
and painted lady (Vanessfe), as well as many others, the sesea 
are alike. This is also the esse with the magnificent Heliconidce, 
and most of the Danaidx in the tropics. But in certain other 
tropical, and in some of our English butterflies, as the 
purple emperor, orange-tip, &c (Apatura Iris and JnthochaTtn 
cariiamines), the sexes differ either greatly or slightly in colour. 
No language suffices to describe the splendour of the males of 

S, p. 123) » peculiar mem- 

observ.iliona, ' The Scottish NbIdi 

ist,' ,Tuly 1672, p. 214. 

Jgs. which is probably con- 

' • The Scottish Naturalist,' J 

fith Hit in-gdui-tion of ths 

1872, p- 313. 

for I.hs iBse of Theoophora, 

• 'Zoological Record,' ISfifi, 


logical Record,' 1869, p. 



Cbaf. XL Butterjlies ana Moths. 309 

Rorae tropical species. Even within the same genus we often 
find speciea presenting estraordinary differences between the 
sexes, wliilst others have their eeses closely ali^a Tbus in the 
South Americoa genus Epicalia, Mr. Bates, to whom I am 
indebted for most of the followii^ facts, and for looking over 
this whole discussion, informs me that he knows twelve species, 
the two sexes of which haunt the same stations (and this ia not 
always the case with butterflies), and which, therefore, cannot 
have been differently affected by esternal conditions." In nine of 
these twelve species the males rank amongst the most brilliant of 
all butterflies, and differ so greatly from the comparatively plain 
females that they were formerly placed in distinct genera. The 
females of these nine species resemble each other in their general 
type of coloration; and they likewise resemble both seses of 
the species in several allied genera, found in various parts of 
the world. Hence we may infer that these nine species, and 
probably all the others of the genus, are descended from an 
ancestral form which was coloured in nearly the same manner. 
In the tenth species the female still retains the same general 
colouring, but the male resembles her, so that he is coloured in 
a much less gaudy and contrasted manner than the males of the 
previous species. In the eleventh and twelfth species, tho 
females depart from the usual type, for they are gaily decorated 
almost like the males, but in a somewhat less degree. Hence in 
these two latter species the bright colours of the males seem to 
have been transferred to the females; whilst in the tenth 
q>ecies the male has either retained or recovered the plain 
colours of the female, as well as of the parent-form of the genus. 
The sexes in these three cases have thus been rendered nearly 
Alike, though in an opposite manner. In the allied genus Eub^is, 
both sexes of some of the species are plain-coloured and nearly 
alike; whilst with the greater number the males are decorated 
with beautiful metalhc tints in a diversified manner, and differ 
much from their females. The females throughaut the genus 
retain the same general style of colouring, so that they resemble 
one another much more closely than they resemble their own 

In the genus Papilio, all the species of the .tineas group 
are remarkable for their conspicuous and stroi^y contrasted 
colours, and they illustrate the frequent tendency to gradation 
in the amount of difference between the sexes. In a few species, 
for instance in F. ascamus, the males and ibmalos are alike; in 

■ See also Mr. BaUs's paper m the same subject, ia regsiil *o 
■Proe. Eat. Soc. of PhitadelphiB,' Diadema, ia ' Transact. Eatomolug. 
Igiio, p. 206. Also Mr, Wallace on Sec. of Loudon,' iee8, p. SJ8. 


310 The Descent of Man. Pabt !L 

others the molefl are either a little brighter, or Tery much mors 
Buperi) than the females. The genus Junonia, allied to our 
Vacesete, offers a nearly parallel coee, for although the sexes uf 
most of the species lesemble each other, and are destitut«i of 
rich colours, yet in certain species, a^ in J. anwue, the male is 
rather more bright-coloured than the female, and in a few (for 
instance J. aiidremiaja) the male is so different from the female 
that he might be mistaken for an entirely distinct species. 

Another striking case was pointed out to me in the Briiiah 
Museum by Mr. A. Butler, namely, one of the tropica,l American 
TheclEe, in which both sesea are nearly aiito and wonderfnCy 
splendid ; in another species the mfile is coloured in a similarly 
gorgeous manner, whilst the whole upper surface of the 
female is of a dull uniform brown. Our common little English 
blue butterflies of the genua Lyciena, illustrate the various dif- 
ferences in colour between the seses, almost as well, though not 
in so striking a manner, as the aboTe exotic genera. In Lycama 
agestis both sexes have wings of a brown colour, bordered with 
small ocellat«d orango spots, and are thus alike. In i. fegon 
the wings of the male are of a fine blue, bordered with black 
whilst those of the female are brown, with a similar border, 
closely resembling the wings of i, agesiis. Lastly,in L. arioa both 
seses are of a blue colour and are very hfce, though ia the femaio 
the edges of the wings are rather duskier, with the black spofa 
plainer ; and in a bright blue Indian speciefi both seses are still 
more alike. 

I have given the foregoing details in order to show, in the first 
place, that when the sexes of butterflies diSer, the male as a 
general rule is the more beautiful, and departs more from the 
usual type of colouring of the group to which the species 
belongs. Hence is most groups the females of the several 
species resemble each other much more closely than do the 
males. In some cases, however, to which I shall herealter 
allude, the females are coloured more splendidly than the 
males. In the second place, thes« details have been given to 
bring clearly before the mind that within the same genus, the 
two sexes frequently present every gradation ftom no difference 
in colour, to so great a difference that it was long before the two 
wore placed by entomologists in the same genus. In the third 
place, we have seen that when the seses nearly resemble each 
other, this appears due either to the male having trausterred 
his colours to the female, or to the male having retained, or 
perhaps recovered, the primordial colours of the group. It also 
deserves notice that in those groups in which tho sexes differ, 
the females usually somewhat resemble the males, so that when 


CttAp, Xi, Butterflies and Moths. 311 

the males are beauidfnl to an extraordinary degree, the females 
almost mvariably exhibit some degree of beauty. Prom tlie 
many cases of gradation in the amount of difference between 
the Bexes, and from the preTalence of the same general type ct 
coloration throughout the whole of the same group, wo may con- 
clude that the causes have generally been the aame which have 
determined, the brilliant colouring of the males alone of some 
epecies, and of both sexes of other species. 

As so many gorgeous butterflies inhabit the tropica, it has 
often been supposed that they owe their colours to the great 
heat and moisture of these zones ; bat Mr. Bates' has shewn by 
the comparison of various closely-allied groups of insects from 
the temperate and tropical regions, that this view cannot be 
ma i n tained ; and the evidence becomes conclusive when bril- 
Uantly-ooloured males and plain-coloured females of the same 
species inhabit the same district, feed on the same food, and 
follow exactly the same habits of life. Even when the sexes 
resemble each other, we can hardly believe that their brilhant 
and beautifully-arranged colours aro the purposeless result of 
the nature of the tissues and of the action of the surrounding 

With animals of all kinds, whenever colour has been modified 
for some special purpose, this has been, as far as we can judge, 
either for direct or indirect protection, or as an attraction between 
the sexes. With many species of butterflies the upper surfaces 
of the wings are obscure; and this in all probability leads 
to their escaping observation and danger. But butterflies 
would be particularly liable to be attacked by their enemies 
when at rest ; and most kinds whilst resting raise their wings 
vertically over their backs, so that the lower surface alone is 
exposed to view. Hence it is this side which is often ooloured 
so as to imitate the objects on which these insects commonly 
rest. Dr. Eiissler, I believe, first noticed the similarity of the 
closed wings of certain Vanessie and other butterflies to the 
bark of trees. Many analogous and striking fa^its could !» 
given. The most interesting one is that recorded by Mr. 
Wallace ' of a common Indian and Stunatran butterfly (Kalliioa), 
which disappears like magic when it settles on a bush ; for it 
hides its head and antennie between its closed wings, which, 
in form, colour and veining, cannot be distingnished &om a 
withered leaf with its footstalk. In some other cases the lower 

• 'The Naturalist oa the Ama- 1867, p. 19. A woodcut of th« 

ions," ToL i. 1863, p. 19. Kallima b given by Mr. Wollaw ib 

' Sw the intraestlng article in ' Hardwicke's Sdenee Qoseip,' Seri, 

lfa« ' Westminilar Review,' July 1SB7, p. 198. 


512 The Descent of Man. Paot IL 

Burfiices of the wings are brillianUy coloured, and jet are 
protective ; thus in Theda rubi the wings when closed are of an 
emerald green, and resemble the young leaves of the bramble, 
on which in spring this butterfly may often be seen seated. It is 
also remarkable that in very many species in which the sexes 
differ greatly in coiom on their upper surfiice, the lower surface 
is closely similar or identical in both eeses, and serves as a 

Although the obscure tints both of the upper and under 
sides of many butterflies no doubt serve to conceal them, yet we 
cannot extend this view to the brilliant and conspicuous colours 
on the upper surface of such species as our admiral and peacock 
Vaneasie, oar white cabbage-butterflies (Pieris), or the great 
swallow-tail Papilio which haunts the open fens — for these 
butterflies are thus rendered visible to every bring creature. 
In these species both seses are alite ; but in the common brim- 
Btone batterfiy ^(ionepteri/a: rhamni), the male is of an intense 
yellow, whilst the female is much paler; and in the orange- 
tip {Anthwharia cardamines) the males alone have their wings 
tipped with bright orai^. Both the males and females in 
these eases are conspicuous, and it is not credible that their 
difiereuco in colour should stand in any relation to ordinary 
protection. Prof. "Weismami remarks,' that the female of one of 
the Lycffinte erpands her brown wings when she settles on 
the ground, and is then almost invisible ; the male, on the other 
hand, as if aware of the danger incurred from the bright blue o( 
the upper surface of his wings, rests with them closed ; and this 
shews that the blue colour cannot be in any way protectiTC 
Neverthelese, it is probable that conspicuous colours are in- 
directly beneficial to many species, as a warning that they are 
unpalatable. For in certain other cases, beanty has been gained 
throagh the imitation of other beautiful species, which inhabit 
the same district and enjoy an immunity from attack by being 
in some way offensive to their enemies; but then we have to 
account for the beauty of the imitated species. 

As Mr. Walsh has remaitod to me, the females of our orange- 
tip butterfly, above referred to, and of an American species 
(AnCi. genutiu) probably shew us the primordial colours of the 
parent-species of the genns; for both seses of four or five 
widely-distributed species are coloured in nearly the same 
manner. As in several previous eases, we may here infer that 
it is the males of Anth. cardamines and genuiia which have 
departed from the usual type of the genus. In the Anth. iara 

• Mr. G. Eraser, In ' Katutf,' • ' EinflusB der Isollrqng anf dk 

Apnl 1871, p. 489. Attbildung,' 1873, p. 58. 


Ci!^. SI. Sutterjlies and Moths. 313 

from Califomia, the orange-tips to the wings have been partially 
developed in the female ; bat the; aie paler than in the male, and 
tlightly different in some other respects. In an allied Indian 
form, the Iphiag glaucippe, the orange-tips are fully developed id 
both sexes. In this Iphias.aa pointed out to me by Mr. A. Butler, 
the under surface of the winga marTeilonsly resembles a pale- 
coloured leaf; and in our English orange-tip, the under surface 
resembles the flower-head of the wild parsley, on whioh the 
butterfly often rests at mght."" The same reason which compels 
us to believe that the lower surfaces ha-re here been coloured for 
the sake of protection, leads us to deny that the wings have 
been tipped with bright orange for the same purpose, especially 
when this charaeter is confined to the males. 

Most Moths rest motionless during the whole or greater part 
of the day with their wings depressed; and the whole upper 
surface is often shaded and coloured in an admirable manner, as 
Mr. Wallace has remarked, for escaping detection. The front- 
wings of the Bombycid» and Noctuidie," when at re^, generally 
overlap and conceal the hind-wings ; so that the latter might be 
brightly coloured without much risk; and they are In fact 
often thus coloured. During llight, moths would often be able 
to escape from their enemies ; nevertheless, as the hind-wings 
are then fully exposed to view, their bright coIoutb muse 
generally have been acc[uired at some little risk. Bat the 
following feet shews how cautious we ought to be in drawing 
conclusions on this head. The common Yeliow Under-wings 
(jCriphaca) often fly about during the day or early evening, and 
are then conspicuous from the colour of their hind-wings. It 
would naturally be thought that this would be a source of 
danger ; but Mr. J. Jenner Weir believes that it actually serves 
them as a means of escape, tor birds strike at these brightly 
coloured and fr^le surfaces, instead of at the body. For in- 
Btanoe, Mr. "Weir turned into his aviary a vigorous specimen of 
Triphtena pronaba, which was instantly pursued by a robin; 
but the bird's attention being caught by the coloured wii^, the 
moth wsjB not captured until after about fifty attempts, and 
small portions of the wings were repeatedly broken oflf. He tried 
the same experiment, in the open air, with a swallow and T. 
fimbria; but the large size of this moth probably interfered 
with its capture." We are thus reminded of a statement made 

"> See the interesting obserTations Science Gossi)!,' Sept. 1S6T, p. 193. 
by Mr. T. W. Wood, ' The Student,' " Sea also, on thb subject, Mr. 

Sept. 18BS, p. 81. Weir's paper in 'Transnct.Enl. Soc. 

" Mt. WallMS in ' Hardwiclia's 1869. p. 23, 


314 The Descent of Man. Pasp 11 

by Mr. Wallace," tjamelj, that in the Brazilian forosts and 
Malayan iRlands, manj common and highly-decorated butterflies 
are weak flyers, though furnished with a broad expanse of wing ; 
and they " are often captnred with pierced and broken wings, 
''as if they had been seized by birds, from which they had 
" escaped : if the wings had been mnch emaHer in proportion 
" to the bodj, it seems probable that the insect would more 
" frequently have been struck or pierced in B vital part, and 
" thus the increased expanse of the wings may have been in- 
" directly beneficial." 

Display. — The bright colonis of many butterflies and of some 
moths are siiecially arrai^d for display, so that they may be 
readily seen. During the night colours are not visible, and 
there can be no doubt that the nocturnal moths, taken as a 
body, are much less ^Wy decorated than butterflies, all of 
which are diurnal in their habits. But the moths of certain 
families, such as the Zygieuidie, several Sphingidce, Uramidee. 
some AratiidiB and Satuiniidje, fly about during the day or 
early evenir^, and many of these are extremely beautiful, being 
far brighter coloured than the strictly nocturnal kinds. A 
few eiceptional cases, Lowever, of brightKioIoQied nocturnal 
species have been recorded.'* 

There is evidence of another kind in regard to display. 
Butterflies, as before remarked, elevate their wings when at 
rest, but whilst basking in the sunshine often alternately raise 
and depress them, thus exposing both surfaces to full view; and 
although the lower surface is often colom^ed in an obscure 
manner as a protection, jet in many species it is as highly 
decorated as the upper surface, and sometimes in a very 
ditTeront manner. In some tropical species the lower surface is 
even more brilliantly coloured than the upper.'* In tre Eng- 
lish fritiUflrieB (^Argynnis) the lower surface »lone is orna- 
mented with shining silver. Nevertheless, as a general rule, 
the upper surface, which is probably more fully exposed, is 
coloured more brightly and diversely than the lower. Hence 
the lower surface generally affords to entomologists the more 

'■ 'Westminster Review,' July Insectsof NewEnglBiid,'lS42,p.315. 
1^67, p. 16. " Such differences betwveu ths 

" For inataoce, Lithosia; bnt upper and lower snrfaces of the 

Prof. Westwood (' Modern Class, of wings of several Bpecios of Papilio, 

Insects,' vol. 11. p. 390) seems Ear- coay bs seen in the beautiful pUte* 

pris«d at this case. On the relative to Mr. Wallace's ' Memoir on the 

colours of diurnal and Docturnal Pspilionidx of the Ualavan Region,' 

Lepidoptera, see ibid. pp. 333 and in 'Transact. Linn. Soc.' toI. iiv. 

S92 ; alio Harris, -Treatise on the part i. 1865. 


Ch*p. XI Butterfiies and Moths. 315 

useful cliaractor for detectii^ tiie affinities cf the Taiious 
Bpecies. Fritz MiiJJer infonns me that three species of Caetnia 
are fbtind near his house ia S, Brazil : of two of them the Iiind- 
wings are obscuie, and are always covered hy the ■frontrwings 
when these hutterffieB are at rest ; but the third spedee tma 
black hiad-wingB, beautifully spotted with red and white, and 
these are fully expanded and displayed whenever the butterfly 
Tests. Other such cases could be added. 

If we now turn to the enormous group of moths, which, as 
I hear from Mr, Stainton, do not habitually expose the under 
surface of their wings to full ■view, we find this side very rarely 
coloured with a brightness greater than, or even equal to, that 
of the upper side. Some exceptions to the rule, either real or 
apparent, must be noticed, as the case of Hypopyra," Mr. 
Trimen informs me that in Guenee's great work, three moths 
are figured, in which the under surface ia much the more 
brilliant. For instance, in the Australian Gastrophora the 
upper surface of the fore-wing is pale greyish-ochieous, while 
the lower svirface is magnificently ornamented by an ocellus of 
cobalt-blue, placed in the midst of a black mark, surrounded 
by orange-yeUow, and this by bluish-white. But the habits of 
these three moths are unsown ; so that no explanation can be 
given of their unusual style of colouring. Mr. Trimen also 
informs me that the lower surfece of the wings in certain other 
Geometrte" and quadrifid Nocture are either more variegated 
or more br^htly-coloured than the tipper surface; but some of 
these species have the habit of " holding their wings quite erect 
" over their backs, retaining them in this position for 3 con- 
" siderable time," and thus exposing the under surface to view. 
Other species, when settled on the ground or herbage, cow and 
then suddenly and slightly lift up their wings. Hence the lower 
surface of the wings being brighter than the upper surface 
in certain moths is not so anomalous as it at first appears. 
The Saturniidie include some of the most beautiful of all 
moths, their wings being decorated, as in our British Emperor 
moth, with fine ocelli; and Mr. T. W. Wood" observes that 
they resemble butterflies in some of their movements; "fc* 
" instance, in the gentle waving up and down of the wings as il 
" for display, which is more chwacteristio of diurnal than of 
" nocturnal Lepidoptera." 

" See Mr. WormaM on this tha Geometre) in 'Traniaet. Eut. 
awth : ' Ptoc. Ent. Soc' March 2nd, Soc' new seiiea, to), v. pi. it. auij 

I (ene of July 6, 1 368, p. i 


3 1 fi -'he Descent of Man. Paet n. 

It is a singular feet that do Britisli moths which are bril- 
liantly coloured, and, as &r as I can discover, hardly any foreign 
Bpeciea, differ much in colour according to sex ; though this ia 
tie case with many briUiaat butterflies. The male, however, oi 
one American moth, the Saiwntia, Jo, is described as having its 
fore-wings deep yellow, curiously marked with purplish-red 
spots ; whilst the wings of the female are purple-brown, marked 
with grey lines." The British moths which differ sexualij in 
colour are all brown, or of various dull yellow tints, or nearly 
white. In several species the males are much darker than the 
females,'" and these belong to groups which generally fly about 
during Uie afteraoon. On the other hand, in many genera, as 
Mr. Stainton informs me, the males have the hind-wings whiter 
than those of the female — of which fact AgroHs exclamationis 
offers a good instance. In the Ghost Moth (NepkUus hamuli) 
the difference is more strongly marked ; the males being white, 
and the females yellow with darker markiiigs.'' It is probable 
that in these cases tie males are thus rendered more conspicuous, 
and more easily seen by the females whilst flying about in 
the dusk. 

From the several forgoing facte it is impossible to admit 
that the briUiant colours of butterflies, and of some few moths, 
h&ve commonly been acquired for the sake of protection. We 
have seen that their colours and eiogant patterns are arranged 
and exhibited aa if for display. Hence I am led to believe that 
the females prefer or are most excited by the more brilUant 

which were fond of eating othei 
moths ; so that if the Cjcnia wits 

. that the mal«s are for the Spilosoma, it would eacapa 
the females in the being devoured, ami iia whilB de- 
_« qaercas, Odanestis poia- ceptive colour would thus be highly 
^^' ^, Hypogymnn dispar, Basyc/ara tenelidal. 
pudibanda, and Cycnia mendiaa. In " It is remnrkable, that in the 
thie latter species the difference in Shetland islands the male of this 
colour between the two seies is moth, instead of differing widely 
strongly marked ; and Mr. Wallace fi-om the female, frequently re- 
informs me that we here have, aa sembles her closely in colour (sea 
he believes, an instance of protective Mr. MacLaehlan, 'Transact. Bnt. 
mimicry confined to one sei, as Sot' vol. ii. IBtiS, p. 459), Mr. 
will hereafter be more fiilly ex- G. Fraser sn^esls (' Nature,' Ariil 
plained. The white female of the 1871, p. 489) that at the season uf 
Cycnia resembles the very common the year when the ghost-moth np- 
Spilotama mentJirasti, hath seies of pears in these northern islands, th* 
which are whiu ; and Mr. Staintcn whiteness of the males wanld nob 
Dhserred that this latter moth was be needed to reudei mem visible ta 
rejected with utter disgust by a the females in the twi'ight night, 
whole brond of young turkeys. 


Cbai-. XI. Butterflies and Mcths. %\^ 

males ; for <nt any other sapposition the males would, as fiii as 
we can see, be ornamented to no purpose. "We know that ants 
and cerfaiia Lamellicom baetlea are capable of feeling an attach- 
ment for each other, and that ants rec<^niao their fellows after 
an interral of several montlis. Hence there is no abstract 
improbability in the Lepidoptera, which probably stand nearly 
or c[nito as high in the scale as these insects, having sufficient 
mental capacity to admire bright colours. They certainly 
discover flowers by colour. The Hamming-bird Sphinx may 
often be seen to swoop down from a distance on a bunch of 
flowers in the midst of green foliage; and I hare been assured 
by two persona abroad, that these moths repeatedly visit flowers 
painted on the walls of a room, and vainly endeavour to insert 
their proboscis into them. Fritz Miiller informs me that several 
kinds of butterflies in S. Brazil shew an unmistakable prefer- 
ence for certain colours over others: he observed that they 
very often visited the brilliant red flowers of five or six genera of 
plants, but never the white or yellow flowering species of the 
same and other genera, growing in the same garden; and i. 
have received other accounts to the same effect. As I hear 
from Mr. Doubleday, the common white bntterfly often flies 
down to a bit of paper on the ground, no doubt mistaking it 
for one of its own species. Mr. Collingwood" in speaking of 
the diifioulty in collecting certain butterflies in the Malay 
Archipelago, states that " a dead specimen pinned upon » 
" conspicuous twig will oft«n arrest an insoet of the same species 
" in its headloi^ flight, and bring it down within easy reach of 
" the net, especially if it be of the opposite sex." 

The courtship of butterflies is, as before remarked, a prolonged 
affair. The males sometunes fight together in rivalry; and 
many may be seen pursuing or crowding round the same 
female. Unless, then, the females prefer one male to another, 
the pairing must be left to mere chance, and this does not 
appear probable. If, on the other hand, the females habitually, 
or even occasionally, prefer the more beautiful males, the colours 
of the latter will have been rendered brighter by degrees, and 
will have beentransmitffid to both sexes or to one sex, according 
to the law of inheritance which has prevailed. The process of 
sexual selection will have been much fitcilitated, if the conclusion 
can be trusted, arrived at irom various kinds of evidence in the 
supplement to the ninth chapter; namely, thai the males of 
many Lepidoptera, at least in the imago state, greatly exceed 
ttie females in number. 

Some facts, however, are opposed to the belief that female 

'' ' Rambles of a Nainralist ia the Chinsae Seas,' 1868, p. 182. 


3l8 The Descent of Man. PisTlI, 

butterflies prefer tto more beautiful males; thus, as I have 
been assured by sDTeral collectors, fresh females may frequently 
be Been paired with battered, faded, or dingy males ; but this is 
a ciTGumstance ivhioli could hardly fail often to follow from tho 
males emerging from their cocoons earlier than the females. 
With moths of the family of the Bomhycidse, the sexes pair 
immediately after assTiming the imago state; for they cannot 
feed, owing io the rudimentary condition of their mouths. The 
females, as several entomologists have remarked to me, lie in an 
almost torpid state, and appear not to evince the least choice in 
regard to their partners. This is the case with the common 
silk-moth (B. mori), as I iave been told by some continental 
and English breeders. Dr. Wallace, who has had great 
esperience in breeding BimAyx cyniMa, is convinced that the 
females evince no choice or preference. He has kept above 
300 of these moths together, and has often found the 
most vigorous females mated with stunted males. The reverse 
appears to occur seldom ; for, as he believes, the more vigorous 
males pass over the weakly females, and are attracted l^ those 
endowed with roost vitality. Nevertheless, the Bombycidaa, 
thoi^h obscurely-coloured, are often beautiful to our eyes from 
their elegant and mottled shades. 

I have as yet only referred to the species in which tho males 
are brighter colonied thaJi the females, and I have attributed 
theii beauty to the feroales for many generations having chosen 
and paired with the more attractive males. But converse 
cases occur, though rarely, in which the females are more 
brilliant than the males ; and here, as I beUeve, the males have 
selected the more beautifiil females, and have thus slowly added 
to their beauty. Wo do not know why in various cksses of 
animals the males of some few species have selected the mora 
beautiful females instead of having gladly accepted any female, 
as seems to be the general mle in the animal kingdom ; but it, 
contrary to what generally occurs with the Iiepidoptera, the 
females were much more numerous than the males, the latter 
would be likely to pick out the more beautiful females. Mr. 
Butler shewed rao several species of CaUidryas in the British 
Museum, in some of which the females equalled, and in others 
greatly surpassed tho males in beauty; for tho females alone 
have the borders of their wings suffused with crimson and 
orange, and spotted with black. The plainer males of these 
species closely rtsemble each other, showing that here the 
females have been modiSed ; whereas in those cases, where the 
males are the more ornate, it is these which have been modified, 
the females remaining closely alike. 


Ohap. XL Butterflies and Motks. 319 

In England we ha^e some analogotis cases, tbongb not eo 
marked. The fenialea alone of two species of Thecla have a 
bright-purple or orange patch on their fore-wings. la Hip- 
parchia the seaes do not difer much; but it is the female oi 
H. janvra which has a conspicuous light-hrowu paich on her 
■wings ; and the females of some of the other species are brighter 
coloured than their males. Again, the females of Coluti edma 
and hyale have " orange or jcllow spots on the blacfe marginal 
" border, represented in the males only by tliin streaks;" and 
in Pieris it is the females which "are oroamented with black 
" spots on the fore-wings, and these are only partially present 
" in the males." Now the males of many butterflies ajre known 
to support the females during their marriage Sight; but in the 
species just named it is the females which support the maJes ; 
so that the part which the two seies play is reversed, as is their 
relative beauty. Throughout the animal kingdom the males 
commonly take the more active share in wooing, and their 
beauty seems to have been increased by the females having 
accepted the more attractive individuals ; but with these but- 
terflies, the females take the more active part in the final mar- 
riage ceremony, so that we may suppose that they Ukewise do 
BO in the wooing; and in this case we can nnderstand how it is 
that they have been rendered the more beautiful. Mr. Meldola, 
from whom the foregoing statements hove been taken, sajs in 
conclusion ; " Though I am not convinced of the action of 
" sexual selection in producing the colours of insects, it cannot 
" l»e denied that these fhots are strildi^ly corroborative of 
" Mr. Darndn's views."™ 

As sexual selection primarily depends on variability, a few 
words must be added on this subject. In respect to colour 
there is no difB.cuIiy, for aay number of highly variable Lepi- 
doptera could be named. One good instance will sufBce. Mr. 
Bates shewed me a whole series of specimens of Fapitia aesostria 
an i P. childrenas ; in the latter the males varied much in the 
extent of the beautifully enamelled green patch on the fore- 
wings, end in the size of tlie white mark, and of the splendid 
crimson stripe on the hind- wings; so that there was a great 
contrast amongst the males between the most and the least 
gaudy. The male of Papilio msostrit is much less beautiful 
than of P, cltildreiKE ; and it likewise varies a little in the size of 

» 'Nntnre,' April 37th, 1871, p. whilst pairing. See alao Mr. G. 

SOS. Mt. Msldola qaotes Donzel, Fraser, in 'Nstnre,' April 2at{i, 

in 'Soc. Ent. de France,' 1S37, p. 1871, p, 489, on the mau differ- 

77, OB the flight of buttei'flin eacas of uieral British tnttsrfliat. 


320 The Descent of Man. Part n 

Ihe green pateii on the fore-wings, and in the occasiona! ap- 
pearance of the Bmall crimson stripe oa the hind-'wings, 
borrowed, as it wonld seem, from its own female ; for the females 
of this and of manj other species in the ^neas group posscaa 
this crimson stripe. Hence between the brightest speoimj'ua 
of P. sesostris end the dnllest of P. childreiiee, there was but a 
small interval ; and it was evident that as far as mere varia- 
bility ja concerned, there wonld be no difficulty in permanently 
increasing the beauty of either species by means of selection. 
The variability is here almost confined to the male sex; but 
Mr. Wallace aod Mr. Bates have shewn" that the females of 
some species are extremely variable, the males being nearly 
constant. In a future chapter I shall have occasion to shew 
that the beautiful eye-like spota, or ocelli, found on the wings of 
many Lcpidoptcra, are eminently variable. I may here add 
that these ocelh offer a difficulty on the theory of sexual 
selection ; for though appearing to ns so ornamental, they are 
never present in one sex and absent in the other, nor do they 
ever differ much in the two sexes." This fact is at present 
inexphcable ; but if it should hereafter be found that the for- 
mation of an ocellus is due to some change in the tissues 
of the wings, for instance, occniring at a very early period oi 
development, we might expect, from what we know of the laws 
of inheritance, that it would be transmitted to both sexes, though 
arising and perfected in one sex alone. 

On the whole, althongh many serious objections may be 
urged, it seems probable that most of the brilliantly coloured 
species of Lepidoptera owe their colours to sesnal selection, 
excepting in certain cases, presently to be mentioned, in which 
conspicuous colours have been gained through mimicry as 
a protection. From the ardour of the male throughout the 
animal kingdom, he is generaOy willing to accept any female; 
and it is the female which usually eserts a choice. Hence, if 
sexual selectiou has been efficient with the Lepidoptera, the 
male, when the sexes differ, ought to be the more brilliantly 
coloured, and this undoubtedly is the case. When both sexes 
are brilliantly coloured and resemble each other, the characters 
acquired by the males apjiear to have been transmitted to both. 

" Wallace on the Papilionidffi of tomolog. Soc" Not. 19th, 1366. p, 

the Malayan Region, in 'Transact, it. 

Liun. Soc.' ya\. iiv. 1865, pp. 8, "■ Mi. Bates trsi so kind m Is 

W. A striking case of a rare lay this subject befots the Eutomn- 

HBTietj, strictly intenoediafe be- logical Society, and I hare receivtJ 

tween two other well-marked female answers to Uiia eflect from MVCTal 


Oflip. XI. Butterflies and Moths. j2i 

We are led fo this conclusion by cases, even within the same 
genuB, of gradation from an extraordinary amoimt of difference 
to identity in colour between tbe two seses. 

But it may be asked whether the differences in colour between 
the seses may not be accounted for by other means besides 
sexual selection. Thus the males and females of the eame 
qwcies of butterfly are in several cases known" to inhabit 
different stations, the former commonly l)aaking in the sunshine, 
the latter haunting gloomy forests. It is therefore possible that 
different oondifiona of lifo may have acted directly on the two 
Beses; bat this is not probable," as in the ada.'t state they are 
exposed to different conditions during a very short period ; and 
the larrte of both are exposed to the same conditions. Mr, 
Wallace believes that the difference between the sexes is due 
not so much to tbe males having been modified, as to the females 
having in all or almost all cases acquired dull colours for tbe 
sake of protection. It seems to me, on the contrary, far more 
probable that it is the males which have been chiefly modified 
through sexnat selection, the females having been comparatively 
little changed. We can thus understand how it is that the 
females of allied species generally resemble one another so much 
more eloselj than do the males. They thus shew us ap- 
proximately the primordial coloTiring of the parent-spcoies of 
the group to which they belong. Thoy have, however, almost 
always been somewhat modified by the transfer to them of some 
of the successive variations, through the accumulation of which 
tbe mates were rendered beautiful. But I do not wish to deny 
that the females alone of some species may have been specially 
modified for protection. In most cases the males and females of 
distinct species will have been exposed during their prolonged 
larval state to different conditions, and may have been thus 
affected ; though with the males any slight change of colour 
thus caused will generally have been masked by the brilliant 
tints gained through sexual selection. When we treat of Birds, 
I shall have to discuss the whole question, as to how far the 
differences in colour between, the sexes are due to the males 
having been modified through sexual selection for ornamental 
pnrposee, or to the females having been modified through 
natural selection for the sake of protection, so that 1 will bere 
say but little on the subject. 

In all the cases in which the more common form of equal 

» H. W. Bates, '.The Natoraliat 
on f-" Amasons,' vol. ii. 1863, p. 
328. A. E, Wallace, in 'Transaet 


322 The Descent of Man. I'art IL 

inheritance hy both sexes has prevailed, the selection of br^ht- 
oolonred malos would tend to mate the females bright-coloured; 
and the selection of dull-coloured females would tend to mate 
the males dull. If both processes were carried oa smmltaneonslj, 
they wonld tend to counteract each other ; and the final result 
wonld depend on whether a greater number of females from 
being well protected by obscure colours, or a greater number of 
males by beii^ brightly-coloured and thus finding partners, 
BUceeeded in leaving more numerous offspring. 

In order to account for the frequent transmission of characters 
to one ses alone, Mr. Wallace expresses his belief tiat the more 
common form of equal inheritance by both seses cau be chansed 
through natural selection into inheritanoa by one sex alone, but 
in favour of this view I can discover no evidence. We know 
from what occurs under domestication that new characters ofteu 
appear, which from the first are transmitted to one sex alone ; 
and by the seleetion of such variations there would not be the 
Blightest difficulty in giving bright colours to the males alone, 
and at the same time or subsequently, dull colours to the females 
alone. In this manner the females of some butterflies and motliB 
have, it is probable, been rendered inconspicuoua for the sake of 
protection, and widely different from their males. 

I am, however, unwilling without distinct evidence to admit 
that two complex processes of selection, eanh requiring the 
transference of now characters to one sex alone, have been 
carried on with a multitude of species, — that the males have 
been rendered mose brilliant by beating their rivals, and the 
females more dull-coloured by having escaped from their 
enemies. The male, for instance, of the common brimstone 
butterfly (Gonepteryx), is of a far more intense yellow than the 
female, though she is equally conspicuous ; and it does not seem 
probable that she specially acquired her pale tints as a protec- 
tion, though it is probable that the male acquired his bright 
colours as a sexual attraction. The female of Antliochans car- 
daminea does not possess the beautiful orange wii^-tij« of the 
male; consequently she closely resembles the white biittorflies 
(Pieris) so common in our gardens ; but wo have no evidence 
that this resemblance is beneficial to her. As, on the other hand, 
she resembles both sexes of several other species of the genus 
inhabiting various quarters of the world, it is probable that she 
has simply retained to a large extent her primordial colours. 

Finally, as we have seen, Tarions considerations lead to the 
(inclusion that with the greater number of brilliantly-coloured 
Iiepidopteca it is the male which has been chiefly modified 
thnnigh Roxual selection; the amount of difference betwees 


Chap. SI. Butterflies and Moths. 32; 

the sexes mostly depending on the fOTm of inheriianoe which 
\ias preyailed. Inheritance is goyemed by bo many nukaown 
laws or conditions, that it seems fo us to act in a eapricjona 
manner ;" and we can thus, to a certain extent, understand how 
it ia that with closely allied species the sexes either differ to an 
astonishing d^ree, or are identical in colour. As all the suc- 
cessive steps in the process or variation are necessarily trans- 
mitted through the female, a greater or less Bumber of sacti 
steps might readily become developed in her ; and thus we can 
understand the frequent gradations from an extreme difference 
to none at all between the sexes of allied species. These cases of 
gradation, it may be added, are much too common to favour the 
auppositioa that we here see females actnally wndei^oing the 
process of transition and losing their hrightmess for the sake of 
protection; for we have every reason to conclude that at any 
one time the greater number of species are in a fixed condition. 

Mimiory. — This principle was fiifit made clear in an admirable 
paper by Mr. Bates,'' who thus threw a flood of light on many 
obscure problems. It had previously been observed that certain 
bntterflios in S. America belongii^ to quite distinct families,, 
resembled the Heliconidte so closely in every stripe and shade of 
colour, that they could not be distinguished save by an ex- 
perienced entomologist. As the Hcliconidte ate coloured in 
their usual manner, whilst the others depart from the usual 
colowing of the groups to whicli they belong, it is clear that 
the latter are the imitatora, and the HeUconidie the imitated. 
Mr. Bates further observed that the imitating species arc com- 
paratively rare, whilst the imitated abound, and that the two 
sets live mingled together. From the fact of the Heliconidie 
beii^ conspicuous and beautiful insects, yet so numerous in 
individuals and species, he concluded that they must be pro- 
tected from the attacks of enemies by some secretion or odour ; 
and this conclusion has now been amply confirmed," especially 
by Mr. Belt. Hence Mr, Bates inferred that the butterflies 
which imitate the protected species have acquired their present 
marvenously deceptive appearance through variation and natural 
selection, in order to be mistaken for the protected kinds, and 
thns to escape being devoured. No explanation is here attempted 
of the hrilhant colours of the imitated, but only of the imitating 
butterflies. We most account for the colours of the former ia 
the same general manner, as in the cases previously discnesed 

" ' The Vsrmiior, of Animak and iiiii. 1862, p. 495. 
Plant; under DoraesticatioD,' vol. ii; " ' Free. Ent. Soa' Dm. 3rd, 


324 "^It^ Descent of Man. Pari II. 

in this chapter. Since the publication of Mr. Bates' paper, 
aimilar and equally striking facts have been obserY&d. by 
Mr. Wallace in the Malayan region, by Mr. Trimen in South 
Africa, and by Mr. Eiley in the United States." 

As some writers have felt much difficulty in underataadin!; 
how the first etepB in the process of mimicry could have been 
effected through natnral selection, it may be well to remark that 
the process probably commenced long ago between forms not 
widely dissimilar in colour. In this case even a slight Tariation 
would be beneficial, if it rendered the one species more like 
the other ; and afterwards the imitated species might bo modi- 
fled to an extreme degree through sexual selection or other 
means, and if the changes were gradual^ the imitators might 
easily be Jed along the same track, until they differed to an 
equally extreme degree from their original condition ; and tbey 
would thus ultimately assume an appearance or oaloiuing wholly 
unlike that of the other members of the group to which thuy 
belonged. It should also be remembered that many species of 
Lepidoptera are liable to considerable and abrupt yariations in 
colour. A few instances have been given in this chapter; and 
many more may be found in the papers of Mr. Bates and 
Mr. Wallace. 

With several species the sexes are alike, and imitate the two 
sexes of another species. But Mr. Trimen gives, in the paper 
already referred to, three cascE in which the sexes of the imitated 
*brm differ from each other in colour, and the sexes of the 
imitating form differ in a like manner. Several cases haTe also 
been recorded where the females alone imitate brilliantlj- 
colonred and protected species, the males retainii^ " the 
" normal aspect of tbeir immediate congeners." It is here obvious 
that the successive variations by which the female has been 
modified have been transmitted to her alone. It is, however, 
probable that some of the many snccessive variations would 
have been transmitted to, and developed in, the males had 
not such males been eliminated by being thus rendered less 
attractive to the females; so that only those variations were 
preserved which were from the first strictly limited in their 
transmission to the female sex. Wo have a partial illus- 
tration of these remarks in a statement by Mr. Eclt;-^' that 

" Wariwn, 'Transact. Linn. Soo.' 163-168. Thia latter essaj- Is vaiu- 

viil.iiv.l865,p. 1 ; alBO ' Transact. abU, ss Mr. Riley hwc discusMs all 

Ent.Ssc' vol. St. (3rd series), 1867, the objentiona which ham been 

p. . 301. Trimen, ' Linn. Transact,' raised gainst Mr. Bates' theory. 
•oL iivi. 1869, p. 497. Riley, " 'Tie Maturaliiit in ^'cuiagns, 

'Third Annual Report on the Noii- 187*, p. 385. 
■na Insects of ttissouri,' 1871, pp. 


vSi-t. X!. Butterflies and Moths. 

Wie males of some of tlie Leptalides, which imitate \ 
species, still rntain ia a concealed manner some of their original 
characters. Thus in the males " the upper half of the lower 
" wing ia of a pure wliitfi, whilst all the rest of the winge ia 
" barred and spotted with black, red and the species 
" they mimic. The females have nfft this whit« patch, and the 
" males nsnally conceal it by coverii^ it with tho upper wing, 
" so that I cannot imagine its being of any other use to them 
" than as an attraction in courtship, when they exhibit it to the 
" females, and thus gratify their deep-seated preference for the 
" normal colour of the Orfer to which the Leptalides belong." 

BrigU Colours of C'lterpillnTS. — Whiht reflecting on the 
beauty of many butterflies, it occunred to me that some cater- 
pillars were splendidly coloured ; and as sesnal selection could 
not possibly have here acted, it appeared rash t» attribute the 
beauty of the mature insect to this af;encj, unless the bright 
colours of their larwe could be somehow explained. In the 
first place, it may be observed that tho colours of caterpillars 
not stand in any close correlation with those of the mature 
insect. Secondly, their bright colours do not serve in any 
ordinary manner as a protection. Mr. Bates informs me, as an 
instance of this, that the most conspicuous caterpillar which he 
ever beheld (that of a Sphinx) lived on the large green leaves of 
a tree on the open llanos of South America; it was about four 
inches in lengUi, transversely banded with black and yellow, 
and with its head, legs, and tail of a bright red. Hence it 
caught the eye of any one who passed by, even at the distance of 
many yards, and no doubt that of every passing bird. 

I then applied to Mr. Wallace, who has an innate genius for 
solving difficulties. After some consideration he replied ; " Most 
" caterpillars require protection, as may be inferred from some 
" kinds being furnished with spines or irritating hairs, and 
" from many being coloured green like the leaves on which they 
" feed, or being curiously like the twigs of the trees on which tJiey 
" live." Another instance of protection, furnished me by Mr. J. 
Mansel Weale, may be added, namely, that there is a caterpillar 
of a moth which lives on the mimosas in South Africa, and 
fobricat«s for itself a case qnit« indistinguishable from the 
surrounding thorns. From such considerations Mr. Wallace 
thought it probable that conspicuously-coloured caterpillars 
were protected by having a nauseous taste; but as their skin 
is extremely tender, and as their intestines readily protrudt 
ftom a wound, a slight peck from the beak of a bird would 
be as iatal to them as if they dad been devcured. Hence, a» 


326 The Descent of Man. Part U 

Mr. ■Wallace rumarks, " distastefulmss alone would be insufBcient 
" to protect a eaterpillar unless Eome outward sign indicated to 
" its would-be destrojer that its prey was a diSKusting morsel." 
Under tbese circumstances it would be higbly advantageous to 
a caterpillar to be instantaneously and certainly recognised as 
unpalatable by aO birds and otber auimalB. Thias the most 
gaudy colours would be serviceable, and might have been ■ 
gained by variation and the survival of the most easily-rc- 
cc-gnised individuals. 

Thia hypothesis appears at first sight very bold, but when it 
was brought before the Entomological Society ^ it was supported 
by various statements ; and Mr. J. Jenner Weir, who keeps a 
large number of birds in an aviary, informs me that he has 
made many trials, and finds no exception to the rule, that ali 
caterpillars of nocturnal and retiring habits with smooth skins, 
all of a green colour, and all which, imitat* twigs, are greedily 
devoured by his birds. The hairy and spinose kinds are 
invariably rejected, as were four conspicuously-coloured species. 
When the birds rejected a caterpillar, they plainly shewed, by 
shaking their heads, and cteansiug their beaks, that they were 
disgusted by the taste," Three conspicuous kinds of cater- 
pillars and moths were also given to some lizards and irogs, by 
Mr. A. Butler, and were rejected, though other kinds were 
eagerly eaten. Thus the probabihty of Mr. Wallace's view is 
Konfirmed, namely, that certain caterpillars have been made 
wnspicuous for their own good, so as to be easily recognised by 
their enemies, on nearly the same principle that poisons are sold 
in coloured bottles by druggists for the good of man. Wo 
cannot, however, at present thus explain the elegant diversity 
in the colours of many caterpOlars ; but any species which had 
at some former period acquired a dull, mottled, or striped appear- 
ance, either in imitation of surrounding objects, or from the 
direct action of cUmate, &c., almost certainly would not become 
uniform in colour, when ite tints were rendered intense and 
bright; for in order to make a caterpillar merely conspicuous, 
there would be no selection in any definite direction. 

Summarj/ and Condudin^ Bemarks on iTtsects. — Looking back 

*' *Proc. Entomolog. Sec.' Dec. analo^us facts in the *Third An- 

3rd, 1866, p. iIt., and Maicli 4th, nual Report on tbs Koi^ns la^c-.t, 

1867, p. Uii. of Missouri,' 1871, p. 148. S<,ma 

"See Mr. J. Jenner Weir's opposed cases ire, however, given bv 

paper DD Insects and Insectivorous Dr. Wallas and M. H. d'Or>ille;'Trausact. Ent.Soc.'lS69, eee 'Zoological Record,' 1869, p. 

p. 21 ; also Mr. Batter's paper, 349, 

ibid. p. 2,1. Mr. Kile; hat give* 


net?. XI. Summary vn hisects. 527 

to the several Ordeie, we see that the seses often differ in 
■various characters, tlie meaning of which is not in the least 
understood. The sexes, also, often differ in their organs of 
Bense and meaiK of locomotion, so that the males may quickly 
discover and reach the females. They differ still oftener in 
the males possessing diversified contrivances for retaining the 
females when found. We are, however, here concerned only in 
a secondary d^roe with sosnal differences of these tinds. 

In almost all the Orders, the males of some epecies, even of 
weak and delicate kinds, are known to be highly pugnacious ; 
and some few are furnished with special weapons for fighting 
with their rivals. But the law of hattle does not prevail nearly 
so widely with insects as with the higher animals. B^ce il 
probably arises, that it is in only a few cases that the males have 
been rendered lai^er and stronger than the females. On the 
contrary, they are usually smaller, so that they may be developed 
within a shorter time, to be ready in large numbers for the 
emergence of the females. 

In two families of the Homoptera and in three of the Orthop- 
tera, the males alone possess sound-prodiiouig oi^ans in an 
eflicient state. These are used incessantly during the broeding- 
Beason, not only for calling the females, bnt apparently for 
oharming or eicitiiig them in rivaby with other males. No 
one who admits the agency of selection of any kind, will, after 
reading the above disonsaion, dispute that these musical instru- 
ments have been acquired throv^ sesual selection. In four 
otler Orders the members of one ses, or more commonly of 
both soses, are provided with organs for producing yarious 
sonnds, which apparently serve merely as call-notes. When 
both sexes are thus provided, the individnals which were able 
to make the loudest or most continuous noise would gain 
partners before those which were less noisy, so that their organs 
have probably been gained through sexual selection. It is 
instructive to reflect on the wonderful diversity of the means 
for producing sound, possessed by the males alone, or by both 
sexes, in no less than six Orders. We thus learn how effectual 
sexual selection has been in leading to modifications which 
sometimes, as with Uie Homoptera, relate to important parts of 
the organisation. 

From the reasons ass^ed in the last chapter, it is probable 
that the great horns possessed by the males of many Lamel- 
licorn, imd some other beetles, have been acquired as ornaments. 
From the small size of insects, we are apt to undervalao their 
appearance. If we conJd imagine a male Chalcoeoma (fig. 16), 
with its polished bronzed coat of maU, and its vast oomjdes 


338 The Descent of Man. Pa^t II. 

norna, magnifiocl to the eizo of a horse, or even of a dog, it would 
be one of the most impOBing aiiimalB in the world. 

The colouring of insects is a complex and ohscure subject 
When the male differs slightlj from, the female, and neither are 
brilliantly-colouied, it is prohable that the Eeses have varied 
in a slightly different manner, and that the variations have been 
transmitted by «ach sex to the same, without any benefit or 
evil thas accruing. When the male is brilliantly-coloured and 
differs conspicviously from the female, as with some dragon-flies 
and many butterflies, it is probable that he owes his colours to 
sexual selection ; whilst the female has retained a primordial or 
very ancient type of colouring, slightly modified by the agencies 
before explained. Eut in some cases the female has apparently 
been made obscure by variations transmitted to her alone, 
as a means of direct protection ; and it is almost certain that 
she has sometimes been made hrilliant, so as to imitate other 
protected species ihhabiting the same district. When tbe sexes 
resemble each other and both ace ot>sourely coloiced, there is 
no doubt that they have been in a multitude of cases so coloured 
for the sake of proteotioiL So it is in some instances when both 
are brightly-coloured, for they thus imitate protected species, or 
resemble surrounding objects such as flowers; or they give 
potice to their enemies that they are unpalatable. In other 
cages in which the sexes resemble each other and are both 
brilliant, especially when the colours are arranged for display, 
we may conclude that they have been gained by the male sex as 
an attraction, and have been transferred to the female. We are 
more especially led to this conclusion whenever the same type 
of coloration prevails throughout a whole group, and we find 
that the males of some species differ widely in colour from 
the females, whilst others differ shghtly or not at all, with 
intermediate gradations connecting these extreme states. 

In the same manner as bright colours have often been 
partially transferred from the males to the females, so it has 
been with the extraordinary horns of many Lamollicom and 
some other beetles. So again, the sound-producing oi^ana 
proper to the males of the Homoptera and Orthoptera have 
generally been transferred in a rudimentary, or even in a nearly 
perfect condition, to the females ; yet not sufficiently perfect to 
be of any use. It is also an interesting fact, as bearing on 
sexual selection, that the stridulatii^ organs of certain male 
Ortbijptera are not fully developed until the last moult ; and that 
the colours of certain male dragon-flies are not fully developed 
until some little time after their emergence from the pupal 
state. Mid when they are ready to breed. 


Obap. XI. Summary on Insects 329 

Sexual BelectioD implies that the more attra<3tiTe mdiTiduols 
tre preferred by the opposite sex; and as with insects, when 
tne sexes differ, it is the male which, with some rare exceptions, 
is tJie more omamentod, and departs moro from the t^pe to 
which the Bpedes helongs ; — and as it is the male which aoarclies 
e^erly for the female, we must supposa that tlie females 
habitually or occasionalUy prefer the more beautiful males, and 
that these haye thus acquired their beauty- That the females 
in most or all the orders would have the power of rejecting 
any particular male, is probable from the many singular oon- 
triTances possessed by the males, such as great jaws, adhesive 
cushions, spines, elongated l^s, Ac, for seizing tho female; for 
these contrivances shew that there is some difficulty in the act, 
80 that her concurrence would seem necessary. Judging from 
what we know of the perceptive powers and affections of 
Taiious insects, there is no antecedent improbability in sexual 
selection having come largely into play ; but we have as yet no 
direct evidence on. this bead, and some facts are opposed to tjie 
belief Nevertheless, when we see many males pursuing the 
same fenmle, we can hardly believe that the pairing is left to 
blind chance — that the female exerts no choice, and is not 
influenced by the gorgeous colours or other omamente with 
which the male is decorated. 

If we admit that the females of tlie Homoptera and Qrthoptcra 
appreciate the musical tones of their male partners, and that the 
Tarions instruments have been perfected through sexual se- 
lection, there is little improbability in the females of other 
insects appreciating beauty in form or colour, and consequently 
in such characters having been thus gained by the males. But 
from the circumstance of colour being so variable, and from it» 
having been so often modified for the sake of protectioa, it is 
difBcuIt to decide in how large a proportion of cases sexual 
selection has played a part. This is more especially difficult in 
those Orders, such as Orthoptera, Hymenopf«ra, and Coleop- 
tera, in which the two sexes rarely differ mnch in colour; for 
we are thon left to mere analc^y. With the Coleoptera, however, 
as before remarked, it is in the great Lamellicom group, placed 
by some authors at the head of the Order, and in which wa 
sometimes seo a mutual attachment between the sexes, that 
we find tho males of some species possessing weapons for sexual 
strife, others famished with wonderful horns, many with stridu- 
lating organs, and others ornamented with splendid metallic 
fjflts. Hence it seems probable that all these characters have 
been gained through the same means, namely sesual selection. 
•With butterflies we have the best evidence, as the males 


no The Descent of Man. Pabt 11, 

sometimes take pains to display their beautifu] colours; and we 
cannot believe that they would act thus, unless the display was 
of use to them in their courtship. 

When wo treat of Birds, we shall seo that they present in 
their Eeoondary seiual charEictors the closest analt^j with 
insects. Thus, many male birds are highly pugnacious, and 
some are furnished with special weapons for fighting with their 
rivals. They possess organs which are used during the hreeding- 
Beason for producing vocal and instrumental miisic. They aro 
frequently ornamented with comhs, horns, wattles and plumes 
of the most diversified kinds, and are decorated with beautifol 
colours, all evidently for the sake of display. We shall find 
that, as with insects, both sexes in certain groups aro equally 
beautifu!, and are equally proyided with ornaments which aro 
usually confined to the male sex. In other groups both sexes 
are equally plain-coloured and unoraamonted. Lastly, in some 
fow anomalous cases, the females are more beautiful than the 
males. We shall often find, in the same group of birds, every 
gradation from no difference between the sexes, to an extreme 
difference. We shall see that female birds, lite female insects, 
often possess more or less plain traces or rudiments of cliaracters 
which properly belong to the males and are of use only to them. 
The analogy, indeed, in all these respects between birds and 
insects is cnrioualy close. Whatever eiplanation applies to the 
one class probably applies to the other; and this explanation, 
as we shall hereafter attempt to shew in further detail, is sexual 

Seoondaet Sexual Chabaotees of Fisheb, Amphibiahb, 


Fishes; Cgnrtshipand Imttles of the tnalea — Larger t\ie of the femi.le* 
— Males, bright colours and omamEntBl appsndages j other itrsngs 
eharaolers — Coloors and appendages acquired bj the maleo durind tha 
breeding-aeasoD alone — Fi^es with both seies brilliaotlj coluuied 
— Proteotive colours— The leas conspicuous colours of the female cannot 

^tAs, and takiDg chai'ge of the ova and ^onng. Aufhibiuis: Dif- 
ferences in strnctufB and colour between the aeiea — Vocal cleans. 
Abp1II,es; Chelonians — CrccodjleB — Suakas, colours in some cases pro- 
tective — Lizarda, battles of— Ornamental appendages — Strange dif- 
fereDces tn structure between the seies — Colours — Seiual diffvrencci 
almost as great as wiSb birds. 


CflAP. SII. Ftshss. 331 

males of Flogiostomous fishes (sharks, rays) and of Cbimteroiij 
fishea are prbTided with claspeis which serve to retain the 
female, like the Tarioaa structures possessed by many of the 
lower animals. Besides tiie claspers, the males of many rays 
have clusters of strong sharp spines on their heads, and seTeral 
rows along "the upper outer surface of their pectoral fins." 
These are present in the males of some species, which hate 
other parts of their bodies smooth. They are only temporarily 
developed during the breeding-season ; and Dr. Giiother suspects 
that they are brought into action as preheasUa oi^;ans by the 
doubling inwards and downwards of the two sides of the body. 
It is a remarkable fact that the females and not the males of 
some species, as of Haia clavata, have their backs studded with 
large hook-formed spines.' 

I'he males alone of the capelin (^MoUotus villosus, one of 
Sahnonidie), are provided with a ridge of closely-set, brush-like 
scales, by the aid of which two males, one on e&cb side, hold the 
female, whilst she runs with great swiftness on the sandy beach, 
and there deposits her spawn.' The widely distinct Mimacanihus 
icopas presents a somewhat analogous structure. The male, as 
Dr. Gunther informs mc, has a cluster of stiff, straight spines, 
like those of a comb, on the sides of the tail; and these in a 
specimen six inches long were nearly one and a half laches in 
length '. the female has in the same pla«e a cluster of bristles, 
which may be comi)ared with those of a tooth-brush. In 
another species, M. peronii, the male has a brush like that 
possessed by the female of the last species, whilst the sides of 
the tail in the female are smooth. In some other species of the 
same genus the tail can bo perceived to be a little roughened in 
the male and perfectly smooth in the female; and lastly in 
others, both sexes have smooth sides. 

The males of many fish flght for the possession of the females. 
Thus the male stickleback (^Oaeterosteta leiurtis) has been de- 
scribed as " mad with deUght," when the female comes out of her 
hiding-pla^e and surveys the uest which he has made for her. 
"He darts round her in every direction, then to his accumulated 
'' materials for the nest, then back again in an instant; and us 
" she does not advance ho endeavours to push her with his snout, 
'.' aod then tries to pull her by the tail aiid side-spine to the nest."' 

' yarrell's ' Hist, of BritUh 1871, p. 119. 
Fishes," vol. ii. 1836, pp. 417, 435, ' See Mr. S. Warington's in- 

thc ipines in S. daoata are peculiar M^. of Nat. Uiat,' Oct. 1852 ml 
ta the female. Not. 1855, 

* ' lite Ameiioaii Katnralist,' April 


332 The Descent of Man. Part !1 

Tho males su« said to be polygamists ;' they are extraotdinarilj 
bold and pugcacious, whiM " the females are quite pacific.'' 
Tlieir battles are at times desperate; "for these puny com- 
" batantfi fasten tight on oach other for several seconds, tumhling 
" over and over agtuu, until their strength appears completely 
" exhausted." With the rough-tailed stickleback (G. tradiuTus) 
the males whilst fighting swim round and loand each other, 
iiitJDg and endeavouiing to pierce each other with their raised 
lateral spines. The same writer adds,' " the bite of these little 
" furies is very severe. They also use their lateral spines with 
" such fatal effect, that I have seen obo during a battle absolutely 
" rip his opponent quite open, so that he sank to the bottom and 
" died." When a fish is conquered, " his gallant beiuing forsakes 
"him; his gay coloura fade away; and he hides his disgraca 
"among his peaceable companions, but is for some time the 
" constant object of his conqueror's persecution." 

The male salmon is as pugnacious as the little stickleback ; 
and so is the male trout, as I hear from Dr. Giiather, Mr. Shaw 
saw a violent contest between two male salmon which lasted 
the whole day ; and Mr. R. Buist, Superintendent of Fisheries, 
informs me that he has often watched from the bridge at Perth 
the males driving away their rivals, whilst the females were 
spawning. The males " are constantly fighting and tearing each 
" other on the spawning-beds, and many so injnre each other as 
" to cause the death of numbers, many being seen swimming near 
" the banks of the river in a state of exhaustion, and apparently 
" in a dying state.'" Mr. Buist informs me, that in June 1868, 
the keeper of the Stormontfield breeding-ponds visited the 
northern Tyne and found about 300 dead salmon, all of which 
with one esception were males ; and he was convinced that they 
had lost their lives by fighting. 

The most curious point about the male salmon is that during 
the breeding-season, besides a slight change in colour, " tho 
" lower jaw elongates, and a cartilaginous projection turns 
" upwards from the point, which, when the jaws are closed, 
" occupies a deep cavity between the intennasillary bones of the 
ipper jaw." ' (Figs. 27 and 2B.) In our sahnon this change of 
structure lasts only during the breeding-season : but in the 

' Ntwl Humphreys, ' River Gar- 


dens,' 1SS7. 

of Salmon Fi: 

ihing,' p. «0) remirln 

• L<mdon'!'MaE.ofNat.HMi.riV 

that like tha 

stag, the male wonid. 

vol. iil. 1830, p. 331. 

if ha could. 

keep all other malu 

' 'Th. Field,' Jane 29tli, 1367. 

For Mr. Shaw's Etntement, sea 


'History of Britiia 

' ErfiDburgh Re.i.«r,' 1843. Anothm 

Fbhes,' vp). ii. 

1836, p. JO. 





Balmo lyi:aodon of N.-W. America the cliange, as Mr. J. K. Lord* 
believes, is permanent, and best marked in the older males which 
have preyiously ascended the riyors. In these old males the 
jaw becomea developed into an immense hook-like projection, ami 

sapermtenijeuce of Dr UDDlinr.] 

the teeth grow into regular fangs, often more than half an iacli 
m lei^h. With the European salmon, according to Mr. Lloyd,' 
the temporary hook-like structure servos 

' Scandinavian Adventures,' vol 


334 The Descent of Man. pAr,T IL 

protect the jaws, Tvh.Gfl one male chains another witli wonderful 
Tiolence ; but the greatly developed teeth of the male American 
salmon may be compared with the tusks of many male mammals, 
and they indicate an offensive rather than a protective pxirpose. 

The salmon is not the only fish in which the teeth differ in 
the two fiesea ; as this is the case with many rays. In the 
thomback (/fnin davata) the adult male has sharp, pointed 
teeth, directed backward^, whilst those of the female are broad 
and flat, and form a pavement ; so that these teeth differ in the 
two sexes of the same species more than is usual in diBficet 
genera of the same family. The teeth of the male become sharp 
only when ho is adult : whilst young they are broad and flat 


Chap. SII. Fishes. 333 

like Uioae of the fetnale. As so frequently oecnrs with secondary 
sexual characters, hoth seses of some speoiea of rajs (for instance 
li. baiis), when adult, possess sharp pointed teeth ; and here a 
character, proper to and primarily gained by the male, appears 
to have been transmitted to the offspring of both sexes. The 
teeth are likewise pointed in both sexes of B. moculata, btrt only 
when quite adult ; the males acquiring them at an earlier ago 
than the females. We shall hereafter meet with analogous 
cases in certain birds, in which the male acquires the plumage 
common to'both sexes when adult, at a somewhat earlier age than 
does the female. With other species of rays the males even when 
old never possess sharp teeth, and consequently the adults of both 
sexes are provided with broad, flat teeth like those of tho yonog, 
and like those of the mature females of the above-mentioned 
species."' As the rays are bold, strong and voraeioiis fish, we 
may suspect that the males require their sharp teeth for lighting 
with their rivals; but as they possess many parts modified and 
adapted for the prehension of the female, it is possible that theii" 
teeth may be used fijr this purpose. 

In regard to size, M. Carbonnier " maintains that tho female of 
almost all fishes is larger than the male ; and Dr. Giinther docs 
not know of a single instance in which the male ie actually 
lai^er than the female. With some Cyprinodonts tho male is 
not even half as large. As in many kinds of fishes the males 
habitually fight together, it is surprising that they haie not 
generally become larger and stroi^r than the females through 
the effects of sexual selection. The males suffer from their 
small size, for according to M. Carbonnier, they are liable to bo 
devoured by the females of their own species when carnivorous, 
and no doubt by other speciea Increased size must be in some 
manner of more importance io the females, than strength and 
siae are to tho males for fightmg with other males ; and this 
perhaps is to allow of the production of a vast number of ova. 

In many species the male alone is ornamented with bright 
colours; or these are much brighter in the male than the 
female. Tho male, also, is sometime'^ provided with appendages 
which appear to be of no mure use to him for the ordinary 
purposes of life, than are the tail feathers to the peacock. I am 
indebted for most of the following facts to the kindness of Dr. 
Giinther. There is reason to suspect that many tropica! fishes 
differ sexually in colour and structure ; and there are some 
strikil^ cases with our British fishes. The male GaEwnymus lyra 

'• S«e Tirrell'i account of the cellcnt fignre. and p. 423. 432. 



The Descent of Man. 


has been called the gemmecus dragonei " from its brilliant gcni- 
" like colours." When fresh caught from the Bea the body is 
yellow of various shades, striped and spotted with vivid blue on 
the head ; the dorsal tins are pale brown with dark longitudinal 
bands; the ventral, caudal, and anal flns being bluish-black. 
The female, or sordid dragonot, was confddercd by Linnjeus, and 
by many subsequent naturalists, as a distinct species ; it is of a 
dingy reddish-brown, with the dorsal fin brown and the other 

fins white. The sexes differ also in the proportional size of the 
head and mouth, and in the position of the eyes ;" but the 
most strikii^ difference is the extraordinary elongation in tbo 
male (fig. 29) of the dorsal flu, Mr. W, Saville Kent remarks 
that this " singular appendage appears from my observations 
" of the species in confinement, to be eubsorvient to tlie same 
" end as the wattles, crests, and other abnormal adjuncts of 
" the male in gallinaceous birds, for the purpose of fascinating 


lish Fishes, 


cajL?. sn. 



" thoir mates."" Tlic young males resemble tho adult femaicB 
in structure and colour. Throughout the genua Callionymus,'* 
the male is generally much more brightly spotted than tha 
female, aad in several species, not only the dorsal, hut the anal 
fin is much elongated in the males. 

The male of the Cottas scorpim, or sea-scorpion, is slenderer 
and smaller than the female, There is also a great difference 
in colour between them. It is difficult, as Mr. Lloyd " remarks, 
" for aoy one, who has not seen this flah during the spawning- 
" season, when its hues are hrighteet, to conceive the admixture 
" of brilliaDt colours with which it, in other respects so ill- 
" favoured, is at that time adorned." Both sexes of the Labrus 
fnixtus, altliOTigh very different in colour, are beautifal; tha 
male being orange with bright blue stripes, and the female 
bright red with some black spots on the back. 

. XipliDphi 

In the very distinct family of the Cyprittodontidte — inhabitants 
of the fresh waters of foreign lands— the sexes sometimes differ 
much in various characters. In the male of the Mollienesta 
pete/tensis," the dorsal fin is greatly developed and is marked 

'5 'NntQi-e,' July 1873, p. 264. " With rsspect to this and the 

^* 'Catnlogneof Acanth. Fishes following species I am indebted to 

tn the British Mnseiim,' by Dr. Dr. Gfiother for information: sea 

Bilnther, 1861, pp. 138-151. also his paper on the 'Fishes of 

" 'Game Birds of Sweden,' Sic, Central America,' in 'Transact. 

1537, p. 486. Zoolog. Soc." vol. vi 1868, p- 485. 


338 Tke Descent of Man. fiW>a 

iviUi a row of Ifo^, round, ocellated. bright- coloured spots; 
n-bilat the same fin in the female is smaller, of ft diflerent shape, 
and marked only witli irregularlj craved brown spots. In the 
male the basal nifli'gin of the anal fin is also a little produced 
iind dark coloured. In the male of an allied form, tte Xipha- 
pharus JJeilerii (fig. 80), the inferior margin of the caudal fin ia 
developed into a long filament, which, as I hear from Dr. Giinther, 
16 striped with hr^t colours. This filament does not contain 
3, and apparently cannot be of any direct use to the 
s in the case of the Callionymus, the males whilst young 
9 the adult females in colour and structure. Sexual 
■differences such as these may be strictly comx)ared with those 
which are so frequent with gallinaceous birds." 

In a siluroid fish, inhabiting the fresh waters of South America, 
the Plecoslomaii harbdtus" (fig. '61), ihsD^le has its mouth and 
inlor-operculiun fringed with a heard of stiff hairs, of wiiich the 
female shows hardly a trace. These hairs are of the nature of 
Bcnlea. In another species of the Ranio genus, soft flexible ten- 
tacles project from the front part of the head of the male, which 
are absent in the female. These tentacles are prolongations of 
the true skill, and therefore are not homologous with the stiff 
hairs of the former species ; but it can hardly be doubted that 
both serve the same purpose. What this purpose may be, it is 
difiicult to conjecture^ ornament does not here seem probable, 
but me can hardly Buppose*that stiff hairs and flexible filaments 
can be usefuJ in any ordinal way to the males alone. In that 
strange monster, the _C/ii™ twit laoiistrosa, the male has a hook- 
shaped bone on the top of the head, directed forwards, with its 
end rounded and covered with sharp spines; in the female" this 
" crown is altf^ther absent," but what its use may be to the 
male is utterly nnknown." 

The structures as yet referred to are permanent in the male 
after he has arrived at maturity ; but with some Blennies, and in 
another allied genus,*' a crest is developed on the head of the 
male only duringthe breeding-season, and the body at the same 
time becomes more brightly-coloured. There can be little loubt 
that this crest serves as a temporary sexual omamimt, fc» the 
female does not exhibit a trace of it. In other species oftthe 
same genus both sexes possess a crest, and in at least one epicies 

" Dr; Gtintter makes this re- Water,' Julj 1868, p. 377, wi* a 

mark ; 'Xktalogaa of Fishis ia the figure. iUay other cases roiilljie 

Britisll lifliseum,' vol. iii. 1861, p. added of structures peculiar to Sie 

-Wl. male, of wliich ths uses are Bill 

V" See Dr. Onnther on tlils Eeniis, known, 

in^a™, 7oolog. Scw.'lSee, p. asi. " Dt, GllntW, 'Catalogue ol 

'■> Bnckiand, in ' Land and Fishes,' vol. iii. pp. 221 and 3«. 


Chap. XII. 'i Fishes. 

Hg. ai. Pbffleloimis bni 


340 The Descent of Man. '.' Pist II, 

neittcr eex is tlniB provided. In many of tho CliroiQidio, for 
instance in Geophagus and especiallj in Ciehia, the males, as 1 
heal from. ProfeESor Agassiz,'" haYe a conspicuous protuberance 
on the forehead, which is wholly wanting in tho feniales and in 
the young males. ProfcEsor Agassiz adds, " I have often 
" observed these fishes at the time of epawning when the pro- 
" tuberance is lai^est, and at other seasons when it is totally 
" wantjng, and the two sexes shew no difference ■whateTer in the 
" outline of the profile of the head. I never conid ascertain that 
" it Bubserves any special function, and the Indians on tho 
" Amazon know nothing about its use." These protuberances 
resemble, in their periodical apyiearance, the fleshy caruncles on 
the heads of certain birds; but whether they serve as omamenti 
must I'emoin at present donbtfol. 

I hear from Professor Agassiz andlir. Giinther.that the males 
of those fishes, which differ permanently in colour from the 
females, oftai become more brilliant during the breeding-season. 
This is likewise the case irith a multitude of fi.ihes, the sexes of 
which are identical in colour at all other seasons of the year. 
The f«nch, roach, and perch may be given as instances. The 
male salmon at this season is "marked on the cheeks with 
" orange-coloured stripes, which give it the appearance of a 
" Labma, and the body partakes of a golden orange tinge. Tlio 
" feniales are dark in colour, and are commonly called bla«k- 
"flsh."** An analogous and even greater change takes place 
with the Salmo eruxE or bull trout; the males of the char 
(S, umUti) are likewise at t-ls season rather brighter in colour 
than the females.'* The colours of the pike ( Esoo: Teticulaiui) of 
liie United States, especially of the male, becotne, durii^ the 
breeding-fieason, exceedingly intense, brilliant, and irideacent." 
Another striking instance out of many is afforded by tho male 
BtJcklebaok (OoBierosiots hivrui), whioh is described by Mr. 
"Warington," as being then " beautiful beyond description." 
The back and eyes of the female are simply brown, and the belly 
white. The eyes of the male, on the othtsr hand, are "of the 
" most splendid green, having a metallic lustre like the groen 
" feathers of some humming-birds. The throat and belly are 
" of a bright crimson, tho back of au ashy-green, and the whole 
" fish appears as though it were somewhat translucent and 

" S8B also ' A Journey in Brazil,' Mag. of Nat. Hiitory,' vol. vi. 184 1, 

by Prof, and Mrs. Agassi!, 1868, p. p. 440. 

220, " -The AniniLan Agdeultiinst, 

" ^amU, 'British Fiahes,' vul. 1868, p. 100. 

IL 1835, pp. 10, 12, 3S. " 'Annalsand Mag. tf Nat. Hist. 

•• W. Thompson, id ■Anaals and Oet, 1852. 


Chat. XII. FMes. 341 

" glowed with an internal incandescence." After the breedii^- 
BffiisoQ these colonts all change, the throat and belly become of a 
paier red, tho back more green, and the glowing tints subside. 

With respect to tho conrtahip of fishes, otJier cases have been 
observed since the first edition of this book appeared, besides that 
ahready given of the stickleback. Mi. W. S. Kent saja that the 
male of the Labru) mio^us, whicb, as we have seen, diflera in 
colour from the female, makes " a deep hollow in the sand of the 
" tank, and then endeavours in tlie most persuasive manner to in- 
" duce a female of the same species to share it with him, swim 
" niiii g backwards and forwards between her and the completed 
" nest, and plainly exhibiting the greatest anxiety for her to follow." 
The males of Cantliarua Uneafus become, diirii^ the breeding- 
season, of deep leaden-black ; they then retire from the shoal, and 
oscaTate a hollow as a nest. "Each male now mounts vigilant 
'' guard over his respectivB hollow, and vigoroiisly attacks and 
" drives away any other fish of the same sex. Towards his com- 
" jmnions of the opposite sex his conduct is far different ; many of 
" the latter are now distended with spawn, and these he endeavours 
" by all the means in his power to lure singly to his prepared 
" LolIow,and there todeposit the myriad ova with whicli theyara 
" laden, which he then protects and guards with the greatest care.'" 

A more striking case of courtship, as well as of display, by tho 
males of a Ghineee Macropus has been given by M. Carbonnier, 
who carefully observed these fishes under confineroent." The 
males are most beautifully coloured, more so tban the females. 
During the breeding-season they contend for tho possession of 
the females ; and, in the act of courtship, expand their fins, 
which are spotted and ornamented with brightly coloured rays, 
in the same manner, according to M. Oarbonnier, as the peacock. 
They then also bound about the females with much vivacity, and 
appear by " I'^talage da leurs vives coulours cherchai a attirer 
" I'atfention des femelles, lesqueUes ne paraissaient indiffcrent«s 
" a ce manfige, elles nageaient avee une molle lentenr vers lea 
" mfiles et semblaient so complaire dans leur voisinage." Aftor 
the male has won his bride, he makes a little disc of froth by 
blowing air and mucus out of his mouth. Ha then collects tho 
fertilised ova, dropped by the female, in his month ; and thia 
caused M. Carbonnier much alarm, as he thought that thoy were 
going to be devoured. But the mala soon deposits them in tho 
disc of froth, afterwards gnardicg them, repairing the froth, and 
taking care of the young when hatched. I mention those par- 
ticolars because, as we shall presently see, there are fishes, the 

■■ ' Satare,' May, 1873, p- 35. 


343 The Descent nf Man Pabt II, 

males of which hatch their cggK in their mouths, and those who 
do not believe in the principle of sradnnl evolution ni^ht ask how 
could BUeh a habit have or^inated , but the difficulty is much 
diminished when we know that tlipre are Bshes which thus 
coUwt and carry the eggs ; for if delayed by any cause in 
depositing them, the habit of hatching them in their moaihs 
might have been aciuired. 

To return to oui more immediate suhjeet. The case stands 
this ; female fishes, as far as I can learn, never willingly spawn 
oicept in the presence of the males ; and the males never fertilise 
the ova except in the presence of the females. The males fight 
for the possession of the females. In many species, the males 
whilst youi^ resemble the females in colour ; but when adult 
become much more briUiant, and retain their colours throughout 
life. In other species the males become brighter than the females 
and otherwise more highly ornamented, only during the season 
of love. The males sedulously court the females, and in one 
case, 88 we have seen, take pains in displaying their beauty 
before them. Can it be believed that thej would thus act to no 
purpose during tiieii courtehip? And this would be the case, 
unless the females exert some choice and select those males 
which please or excite them most. If the female exerts such 
choice, all the above facts on tte ornamentation of the males 
become at once intelligible by the aid of sexual selection. 

"We have next to enquire whether this view of the bright 
colours of certain male fishes having been acquired through 
aesual selection can, through the law of the equal transmission of 
characters to both sexes, be extended to those grouje in which the 
toaleB and females are brilliant in the same, or nearly the same 
degree and manner. In such a genus as liabrus, which includes 
some of the most splendid fishes in the world — for instance, the 
Peacock Labms {L. pavo), desoribed,*' with pardonable exaggera- 
tion, as formed of polished scales of gold, encrusting lapia-lazuli, 
rnbies, sapphires, emeralds, and amethyBts— we may, with much 
probability, accept this belief; for we have seen that the eexoe in 
at least one species of the genus differ greatly in colour. With 
some fishes, as with many of the lowest animals, splendid colours 
may be the direct result of the nature of their tissues and of the 
surroimding conditions, without the aid of selection of any kind. 
The gold-fish (fiyprinus auratas), judging from the analogy of 
the golden variety of the common carp, is perhaps a case in point, 
as it may owe its splendid colours to a single abrupt variation, 
due to the conditions to which this fish has been subjected nnder 

» Bory de Siunt Vincent, in ' Diet. ClaiB. dllist. Kal.' tem. ii. tSiS 


Cbat. XII. Fisfies. 343 

oonflnaiient. It is, however, more probable ttat these coloitra 
nave been intensified through artificial selection, aa this speeioa 
fias been carefully bred in China from a remoi« period." Under 
natural conditions it does not eeem probable tbat beings so 
Dighly organised aa fishes, and which live under snoh complex 
relations, should become brilliantly colonred without sufiering 
norne evil or receiving some benefit from so great a change, and 
consoqTiently without the intervention of natural selection. 

WTiat, then, are we to conclude in regard to the many fishes, 
both sexes of which are splendidly coloured? Mr. Wallace" 
beheves that the species whioli frequent reefs, where corala and 
other brightly -coloured organisms abound, are brightly coloured 
in order to escape detection by their enemies; but according to 
. my recollection they were thus rendered highly conspicuous. 
In the fresh-waters of the tropics ■ there are no brilliantly- 
coloured corals or other organisms for the fishes to resemble ; 
yet many species in the Amazons are beautifully coloured, and 
many of the carnivorous Cyprinidte in India are ornamented 
with "bright longitudinal lines of various tints."*' Mr. M'Clel- 
land, in describing these fishes, goes so far as to suppose that 
" the peculiar briHianey of their colours " serves as " a better 
" mark for king-fishers, terns, and other buds which are 
" destined to keep the number of these fishes in check ; " but at 
the present day few naturalists will admit that any animal has 
been made conspicuous aa an aid to its own destruction. It is 
possible that certain fishes may have been rendered conspicuous 
in order to warn birds and beasts of prey that they were 
unpalatable, as explained when treating of caterpillars ; but it 
is not, I believe, known that any fish, at least any fresh-water 
fish, is rejected from being distastefal to fish-devouring animals. 
On the whole, the most probable view in regard to the fishes, of 
which both sexes are brilliantly coloured, is that their colours 
were acquired by the males i 
transferred equally or nearly si 

** g mm has cen " prodnced at Hangohow a 

(d mad m w On riety culled the fire-fish, from its 

f m Dm enaely red colour, it is uni- 

W Esllj admired, and there h not, 

N Q es, A 6 ousehold where it is not cul- 

pe H as a source of profit." 

g w rs re 'Westminster Rerietr," Julj 

00 fi m g Son Dy 8 p. 7. 

S3 comoi cd ■ ladian Cypiinidas,' bj Mr. 

ear fi es M elland, 'A-iiatic RpRearckfo, 

ed la tm p ac ii. part ii. 1839, p. 230. 


The Descent of Man. Past IL 

We liaTe now to consider whether, when the male differs in a 
marked manner from the female in colonr or in other orna- 
ments, he alone haa been modified, the Tariations being inherited 
by his male ofepring alone; or whether the female has been 
epecially modified and rendered inconspionous for the sake of 
protection, such modifications beii^ inherited only by the 
females. It is impossible to donbt that colour has been gained 
by many fishes as a protection : no one can examine the speckled 
upper snrfece of a flounder, and OTerlook its resemblance t« the 
sandy bed of the sea on which it lives. Certain fishes, moreover, 
can through the action of the nervous system, change their 
colours in adaptation to snirounding objects, and that within a 
short time."" One of the most striking instances ever recorded 
of an animal being protected by its colour (as far sa it can bo 
judged of in preserved specimens), as well as by its form, is that 
given by Dr. Giinther" of a pipe-fish, which, with its reddish 
streaming filaments, is hardly distinguishable from the sea-weed 
to which it clings with its prehensile tail. But the question now 
under consideration la whether the females alono have been 
modified for this object. We can see that one ses will not be 
modified throngh natural selection for the sake of protection 
more than the other, suppodcg both to vary, unless one sex is 
exposed for a longer period to danger, or has less power of 
escaping from such danger than the other; and it does not 
appear that with fishes the sexes differ in these respects. As 
far as there is any difference, the males, from being generally 
smaller and from wandering more about, are exposed to greater 
danger than the females; and yet, when the sexes differ, the 
males are almost always the more conspicuously coloured. 
The ova are fertilised immediately after being deposited; and 
when this process lasts for several days, as in the case of 
the salmon," the female, during the whole time, is attended by 
the mal& After the ova are fertilised they are, in most casce, 
left unprofeoted by both parents, so that the males and females, 
as fiir as oviposition is concerned, are equally exposed to danger, 
and both are equaOy important for the production of fertile ova ; 
consequently the more or less brightlr-coloured individuals of 
either sex wonld he equally liable to be destroyed or preserved, 
rrad both wonld have an equal influence on the colours of their 

Certain fishes, belonging to several families, make nests, and 
some of them take care of tteir young when hatched. Both 

" G. Ponehet, L'Institat. Nov. 1, 327, pi. liv. and it. 
;9T1, p. 13i. " Yarrell, ' British FUkei,' voi 

« •?!«:. Ziwlog. Soc." 1866, p. iL [.. 'I 


Cuke. xn. Fis^s. 34S 

Beses of the br^ht eoIoTiied Crenilabrus massa and melons 'wori 
together in building their neste with sea-weed, etiells, Ac." 
But the niales of certain fishea do all the work, and aftertrcirda 
tato osclnBiTO charge of the young. This is the case with the dull- 
coloured gobies," in which the eeies are not kuown to differ in 
oolonr, and likewise with the sticklebacks (Gasterosteus), in which 
the males become briUiantl; coloured during the spawnii^ season. 
Tte mate of the smooth-tailed stickleback (0. leiurus) performs 
the duties of a nurse with exemplary care and valance daring 
a long time, and is continually employed in gently leading back 
the young to tte nest, wlien they stray too fitr. He courageoufily 
drives nway all enemies, including the females of his own species. 
/t would indeed be no small relief to the male, if the female, after 
depositing her ^g^, were immediately devoured by some enemy, 
for he is forced incessantly to drive her from the nest^" 

The males of certaia other fishes inhabiting South America 
and Ceylon, belonging to two distiaet Orders, have the estra- 
ordioary habit of hatching within their mouths or branchial 
cavities, the e^s laid by the females." I am informed by 
Professor Agassiz that the males of the Amazonian species 
wliich follow this habit, " not only are generally brighter than 
" the females, but the difference is greater at the spawning-season 
" than at any other time." The species of Geophagns act in the 
same manner; and m this genus, a conspicuous protuberance 
becomes developed on the forehead of the males during the 
breeding-season. With the various species of Ghromids, as 
Professor AgaBsiz likewise informs me, seiuol differences in 
colour may be observed, " whether they lay their eggs in the 
" water among aquatic plants, or deposit them m holes, leaving 
" them to come out without further care, or build shaDow nests 
" in the river mud, over which they sit, as our Pomotis does. 
" It ought also to be observed that these sitters are among the 
" brightest species in their respective families ; for instance, 
" Hygrogonus is bright green, with large black ocelli, encircled 
" with the most briUiant red." Whether with all the species of 
Chromids it is the male alone which sits on the eggs is not 
known. It is, however, manifest that the fact of the e^s being 

■' AccocdioE to the gliservitions naU andMng. of Nat. Hist.' Novem- 
of M. Gerbe ; see GSnther's ' Record ber 1855. 
of Zooiog. Literatore,' I8S5, p. " Prof. Wyman. in 'Pros. Boston 

" Covier, 'K6gne Animal,' vol. Also Prof. Tamtt, id 'Joarnal of 

J. 1829, p. 242. Anatomy and Php.' Nov. 1, IMS, 

^„ ,, -^ .. ..... . p^g jjj. Gdnther haa likesiB* 


J46 Th£ Descent of Man. Pasi TI. 

protected or nBprotected b; the parents, has had h'ttle or no 
influence on the differences in colour between the sexoe. It ia 
further raanifest, iu all the cases in which the males take 
eiclnsive charge of the nests and young, that the destruction 
of the biighter-eoloured males would be fai more influential on 
the character of the race, than the destruction of the brighter- 
coloured females; for the death of the male during the period of 
incubation or narsing would entail the death of the young, eo 
that they could not inherit hia peculiarities; jet, in many of 
these very cases the males are more conspicuonfily coloured than 
the females. 

Inmost of the Lophohranchii (Pipe-fish, Hippocampi, &c.) 
the males have either marsupial sacks or hemispherical de- 
pressionR on the abdomen, iu which the ova laid by the female 
are hatched. The males also shew great attachment to their 
young." The sexes do not commonly differ much in colour; 
but Dr. Giinther believes that the male Hippocampi are rather 
brighter than, the females. The genus Solenostoma, however, 
offers a curionB exceptional case,*' for the female is much more 
vividly-coloured and spotted than the male, and she alone has a 
marsupial sack and hatches the eggs; so that the female of 
Solenostoma differs from all the other Lopbot>ranchii in this 
latter respect, and from almost all other fishes, in being more 
brightly-coloured thau the male. It is improbable that this 
remarkable double inversion of character in the female should 
be an accidental coincidence. As the males of several fishes, 
which take eiclnsive charge of the e^s and young, are more 
brightly coloured than the females, and aa bore the female Sole- 
nostoma takes the same charge and is brighter than the male, it 
might be argued that the conspicuous colours of that ses which 
is the more important of the two for the welfare of the ofepring, 
must be in some manner protective. But from the large number 
of fishes, of which the males are either permanently or period- 
ically brighter than the females, but whose life is not at all 
more important for the welfare of the species than that of the 
female, this view can hardly be maintained. When wo treat 
(rf birds we shall meet with analc^ous cases, where tbere has 
been a complete inversion, of the nsnal attributes of the two 
sexes, and we shall then give what appears to be the probable 
explanation, namely, that the males have selected the more 
attTftctJve females, instead of the latter having selected, ia 

»• TarTell, 'Hist, of Britieh Fhhes of Zaniihar,' by Col. Plsffsir, 

FishBs,' ml. ii. 1836, pp. 329, 338. 1B66, p. 137, h.Ts re-iiamined tb. 

" Dr. OUnthsr, liace pDblisbiDg specim^iis, and has givau au the 

in KcooEt of this species in 'The aJjovB inforniHtion. 


Chap. XII. FisJus 347 

eccordance with, the TKnal rule throoghout the animal kingdom, 
the more attractiye males. 

Oa tie whole we may conclude, that with moat fishes, in 
which the seies differ in colour or in other ornamental charac- 
ters, the males originallj Taried, with thoir Tariationa trans- 
mitted to the same sex, and acciomnlatod through sexual 
Belection by attracting or exciting the females. In many cases, 
LoweTer, such characters have been transferred, either partiaDy 
or completely, to the females. In other cases, again, both seies 
have been coloured alike for the sake of proteetioii; but in 
no instance does it appear that the female alone has had her 
colonra or other characters spocially modified for this latter 

The last point which need be noticed is that fishes are known 
to make Tarious noises, some of which are described as being 
musicaL Dr. Dufoss^, who has especially attended to this 
subject, says that the sounds are volantarily produced in several 
ways by different fishes : by the friction of the pharyngeal bones 
—by lie Tibration of certain muBcles attached to the swim- 
bladder, which Beryes as a resounding board — and by the vibra- 
tion of tbe intrinsic muscles of the swim-bladder. By this latter 
means the Trigla produces pure and long-drawn sounds which 
range over nearly an octave. But the most interesting case for 
US is that of two species of Ophidium, in which the males alone 
are provided with a sound-producing appaiatus, consisting of 
small movable bones, with proper muscles, in comiection with 
tha swim-bladder." The drumming of the TJmbrinas in the 
European seas is said to be audible from a depth of twenty 
fathoms ; and the fishermen of Bochelle assert " that the males 
" alone make the noise during the spawning-time ; and that it 
" is possible by imitating it, to take them without bait'"' From 
this statement, and more especially from the case of Ophidium, 
it is almost certain that in this, ttie lowest class of the Verte- 
brata, as with so many insects and spiders, Bonnd-produoing 
instruments have, at least in some cases, been developed through 
sexual selection, as a means for bringing the sexes together. 

•I ' Comptes Rendus." Tom. ilrt. the Dutrh trnBslation of this work 

1858, p. 353. Tom. iWii. 1858, p. (vol. ii., p. 38), gives some further 

816. Tom. lir. 1862, p. 393. The particulars oo th« sounds made by 

noiBu luade bythe Umbrinas(S!»(cna fishes. 

oguaaX l» sail! b? some authors to " The Rev. C. Kingsley, in 

bemoraliltBthatof aflote DTorgan, 'Natnie,' May 1870, p. 40. 
tbu drtuuming : Dr. KouUTten, is 


The Descent of Man. 


Urode^a. — I will begin with the tailed -imphihians Tlie fcxea 
of salamanders or newts often diffi,r much hoth m colour ind 
gttucture. In some species jreliensile clawi are developed on 
the fore-legs of tlie males dtmng tlie breedmg season and at 
this season in the male Ti-iton ptlmipei the limd feet ire pro 
Tided with a swimming-web whieh ir almost completely 
absorbed during the winter; so that their fict then rL'^mlle 

f ■ ""^N^ 

those of the female." This structure no doubt aids tlio male 
in his eager search and pursuit of the female. "Whilst courting 
her he rapidly Tibratrai the end of his tail. With our common 
newts (Triton punctatits and cristatm) a deep, much indented 
crest is developed along the back and tail of the male during lie 
breeding-season, which disapiiears during the winter. Mr. St. 
Geoige Mivart informs me that it is not furnished with muscles, 
and therefore cannot be used for locomotion. As during the 
ceason of courtship it becomes edged with bright colours, there 
can hardly he a doubt that it is a masculine ornament. In 
many species the body presents strongly contrasted, though 
lurid tinte, and these become more Tivid during the tirooding- 
season. The male, for instance, of our common little newt 
(Tritim punctot'is) is " brownisli-grey above, passing into yellow 

' Bell, ' History of Britisli Reptiles,' 2iid sdit. 1849, pp. 156-1 


Chap. IIL Amphibians. 349 

" beneath, which in the spring becomes a rich briglit orange, 
" njHjKed everywhere with, round dark spots." The edge of the 
crest also is then tipped with bright red or Tiolet The female 
is usually of a yellowish-brown colom with scattered brown 
dots, and the lower surfiico is often quite plain." The young 
are obscurely tinted. The ova are fertilised during the act of 
deposition, and are not subsequently tended by either parenf. 
We roay therefore conclude that the males have acquired their 
strongly-marked colours and ornamental appendages through 
sexual selection ; these being transmitted either to the male 
of&pring alone, or to both seies. 

AnuTo. or Batraokia. — With many frc^ and loads the colours 
evidently serre as a protection, such as the bright green tints 
of ttee-frogs and the obscure mottled shades of many terrestrial 
species. The most conspicuously-coloured toad which I ever 
saw, the PhryntFeus ntgrUtins,"' had the whole upper surface of 
the body as hhxk as ink, with the soles of the feet and parts of 
the abdomen spotted with the brightest vermilion. It crawled 
about the bare sandy or open grassy plains of La Plata under a 
scorcliing sun, and could not fail to catch the eye of every pass- 
ing creature. These colotirs are probably beneficial by making 
this animal known to all birds of prey as a nauseous mouthful. 

In Nicaragua there is a htUe irog " dressed in a bright livery 
" of red and blue " which does not conceal itself like most other 
species, but hops about during the daytime, and Mr. Belt says" 
that as soon as he saw its happy sense of security, he felt sure 
that it was uneatable. After several trials he succeeded in 
temptii^ a young duck to snatch up a young one, but it was 
instantly rejected; and the duck "went about jerking its head, 
" as if trying to throw off some unpleasant taste." 

With respect to sexual differences of colour, Dr. Giinther 
does not know of any striking instance either with fi'ogs or 
toads ; yet he can often distinguish the male from the female, by 
the tints of the former being a little more intense. Kor does 
he know of any striking difference in external structure between 
the sexes, excepting the prominences which become developed 
during the breeding-season on the iront^Iegs of the male, by 
which he is enabled to hold the female." It is surpriaing thai 

" Bell, ' HJFlory of British liep- 
tiien,' 2nd edit. 1849, pp. 146, 151. 

" 'Zoology of the Voyag* of llie piate-llliB callosities on the thumi 

" Beagle," ' 1S43. Bell, ibid. p. 49. and cortsia rugosities nn the hugers, 

" ' The tiaturalist in NicBtagus,' which perhaps snb&erve the same an^ 


350 The Descent of Man. Pabt R, 

theso ammala ta^e not acciuired more sttongly-markcd sexaal 
clmracteis ; for thoagli cold-blooded their passions are stronfs 
Dr. Giinther intonna me tliat he has several times found oh 
■nnfortiwiate female toad dead and Bmothered from ha^ng been 
so closely embraced by three or four males. Frogs have been 
observed by Professor Hoffman in G-iessen fighting all day long 
during the breeding-season, and with so much violence^ that one 
iiad its body ripped open. 

Frt^^ and toads offerone interesting sexual difierence, namely, 
in the musical powers possessed by the males; but to speulc 
of music, when applied to the discordant and overwhelming 
sounds emitted by male bull-frogs and some other species, seems, 
accordit^ to our taste, a singularly irappropriate expression. 
Nevertheless, certain fn^s sing in a decidedly pleasing manner. 
Near Eio Janeiro I used often to sit in the evening to listen to a 
number of little Hylie, perched on blades of grass close to the 
water, which sent forth sweet chirping notes in harmony. The 
various sounds are emitted chiefly by the males during thn 
breeding-season, as in the case of the croaking of our common 
frc^.'* In accordance with this fact the vocal ot^ns of the 
males are more highly-developed than those of the females. In 
some gencm the males alone are provided with sacs which open 
into the larynx." For instance, in tte edible frog (liana escuknta) 
" the saes ore peenliar tc the males, and become, when filled 
" with air in the act of croaking, large globular bladders, stand- 
" ing out one on each side of the head, near the corners of the 
•' mouth." The croak of the male is thus rendered exceedingly 
powerful ; whilst that of the female is only a slight Kroaning 
noise." In the several genera of the family the vocal organs 
differ consiiierably in structure, and tlieir development in ail 
cases may bo attributed to sexual selection 


Cheloiiia. — Tortoises and turtles do not offer well-marlicd 
sexual differences. In some species, the tail of the male is 
longer than that of the female. In some, the plastron or lower 
Burfaee of the shell of the male is slightly concave in relation lo 
the back of the female. The male of the mud-t«rtle of the 
United States (Chrgsemyt picia) has claws on its front-feet 
twice as long as those ot the female ; and these are used when 

'• Bell, ' Histerj of British " J. Bishop, in ' Todd'e Cyclop. 

Reptiles,' 1849, p. 93. of Anat. md Phvs.' vol. iv. p. 1503 

•0 Ball, ibid. p. 112-114, 


CaAl>. XJI. Reptilei. %t,\ 

the sexes umto." 'Witti the huge tortoiso of tbo Gaiapago* 
Islands (Testudo nigra) the mali« are said io grow to a larger 
Bize than the females; during the pairmg-season, and at eo 
other time, the male utters a hoarse bellowing noise, which can 
be heard at the distance of more than a hundred yards ; the 
female, on the other hand, nerer uses her Toiee." 

With the Tuiliidv elegans of India, it is said " that the eombata 
" of the males may be heard at some distance, from the noise 
" they produce in butting against each other."" 

Crocociffia.— The seses apparently do not differ in colour ; nor 
do I know that the males fight together, though ttiis is pro- 
bable, for some kinds mate a prodigious disphiy before the 
females. Bartram" describes the male alligator as striTicg 
to win the female by splashing and roaring in the midst 
of a lagooo, *' swollen to an extent ready to burst, with its 
" head and tail lifted up, he spins or twirls round on the 
" surface of the water, like an Indian chief rehearsing his feats 
of war." During the season of Ioto, a musky odour is emitted 
by the Eubmasillary glands of the crocodile, and pertadcs their 

Ojihidia. — Dr. Giinther informs me that the males are always 
smaller tlian the females, and generally have longer and slenderer 
tails; but he knows of no other difference in external stmotnre. 
In regard to colour, he can almost always distinguish the male 
from the female by his more strongly-pronounced tints ; thus 
the black zigzag band on the back of the male English viper is 
more distinctly defined than in the female. The difference is 
much plainer in the rattle-snakes of N. America, the male of 
which, as the keeper in the Zoological Gardens shewedmc, can at 
onee be distinguished from the female by having more lurid 
yellow about its whole body. In S. Africa the JiiKep/nUna 
capensis presents an anali^ous difference, for the female "is 
" never so fully variegated with yellow on the sides as the 
"male."" The male of the Indian Hipsas cynodoa, on the 
other hand, is blackish-brown, with the belly partly black, 
whilst the female is reddish or yellowish-olive, ivith the belly 
either uniform yellowish or maibled with hlaek. In the Tragops 
dispar of the same country, the male is bright green, and the 

"Mr. a J. Majrnard, "The British India,' 1804, p. 7, 
AmsriaiB Naluniliet,' Due 1869, p. " "Pi'avsin throngh Carulina, 

65.V *K., 17W1, p. 128, 

" Soe ray 'Jonrosl of Eeaearches " Ow*n, ' Anatomy of Verle- 

duriog the Vojageofthe" Beagle,"' bratoa,' vol. i. 1866, i>. 615. 
184&, p. S84. 


352 The Descent of Man. Paut H 

female bronze-coloured." No doubt the colours of soma anakea 
aw protective, as etemi by the green tints of tree-snakes, and 
the YariouB mottled shadoa of tie species which live in sandy 
places ; bnt it is doubtful whether tbe colours of many kinds, 
for instance of the common Ei^lish snake and viper, serve to 
conceal thein ; and this is still more doubtful with the many 
foreign si)eoiea which are coloured with estreme elegance. The 
colours of oertaia species are very different in the adult and 
young BtateK.** 

During the breeding-season the anal scent-glands of snakes are 
in active function ;" and so it is with the same glands in lizards, 
and as we have seen with the submaxillary glands of crocodiles. 
As the males of roost animals search for the females, theao 
odoriferons glands probably serve to excite or charm the fen ale, 
rather than to guide her to the spot where the male may ba 
found. Male snakes, though appearing so sluggish, are amorous ; 
for many have been obserscd crowding roimd the same female, 
and even round her dead body. They are not known to 
fight tc^ther from rivalry. Their intellectual powers are 
higher than might have been anticipated. la the Zoological 
(Jardens they soon leam not to strike at the iron bar with which 
their cages are cleaned ; and Dr. Keen of Philadelphia informs 
me that some snakes which he kept, learned after four or five 
times to avoid a noose, with which they were at first easily 
caught. An excellent observer in Ceylon, Mr. E. Layard, saw " 
a cobra thrust its head through a narrow hole and swallow a 
toad. " With this encumbrance he eould not withdraw him- 
" self; finding this, he reluctantly disgorged the precious mor- 
" sel, which began to move off; this was too much for Hcake 
" philosophy to bear, and the toad was again seizedj and again 
" was the snake, after violent efforts to escape, compelled to part 
" with its prey. This time, however, a lesson had been learnt, 
" and the load was seized by one leg, vrithdrawn, and then 
" swallowed in triumph." 

The keeper in the Zoological Gardens is positive that certain 
Bnakes, for instance Crotalus and Python, distii^uish bim from 
all other persons. Cobras kept together in the same ca^T 
apparently feel some attachment towards each other.'* 

" Dr. A. QUnthet, 'Rrptiles of braWs," vol. i. 1866, p. 615. 
Britifih India,' Ray Sot 1864, pp. '" 'RanibleaiiiCsylon,'iii'AlLD!lil 

304, 308. aod Mag. of Nat. Hist.' 2nd lene., 


Ohaf. xn. Reptiles. 353 

It does not, however, follow because snakes have soma 
reasonii^ power, strong passions and matual affection, that they 
ehould likewise bo endowed with sufEeiiint, taste to admire 
brilliant colours in their partners, so as to lead to the adorn- 
ment of the species through sesual selection. NeTerthelesa, it is 
difficult to account in any other manner for the estreme tcauty 
of certain species ; for instance, of the coral-snakes of S. America, 
which are of a rich red ivith black and yellow transverse bands. 
I well remember how much surprise I fe!t at the beauty of the 
first coral-snake which I saw gliding across a path in Brazil. 
Snakes coloured in this peculiar maoner, as Mr. Wallace states 
on the autJiority of Pr, Giinther,'^ are found nowhere else 
in tiie world except in S. America, and here no less than four 
genera occur. One of those, Slaps, is venomous ; a second and 
widely-distinct genus is doubtl'ully venomous, and the two 
others are quite harmless. The xpecies belonging \a these 
distinct genera inhabit the same districts, and axe so like each 
other, that no one "but a naturalist would distinguish the 
" harmless from the poisonous kinds," Hence, as Mr. Wallace 
believes, the innocuous kinds have probably acquired their 
colours as a protection, on the principle of imifation ; for they 
would nattttally be thought dangerous by their enemies. The 
cause, however, of the bright colours of the venomous Elai>s 
remains to be explained, and this may perhaps be sexual 

Snakes produce other sounds besides hisaii^. The deadly 
Eiiliia mrmata has on its sides some oblique rows of scales of a 
peculiar structure with serrated edges ; and when this snake is 
excited, these scales are rubbed against each other, which pro- 
duces " a curious prolonged, aJmost hissing sound." ™ With 
respect to the rattling of the rattle-snake, we have at last some 
definite information : for Profissor Anghey slates," that on twc 
occasions, being himself unseen, he watched from a little distance, 
a rattle-snake coiled up with head erect, which continued to 
rattle at short intervals for half an hour ; and at last he saw 
another snake approach, and when they mot they paired. 
Hence he is satisfied that one of the uses of the rattle is to bring 
the sexes together. Unfortunately he did not ascortain whether 
it was the male or the female which remained stationary and 
called for the other. But it by no means follows from tha 
above fiict that the rattle may not be of use to these snakes in 
ather ways, as a warning to animals which would otherwise 

« ' Westminater Revie*,'Jnli 1st, S"c' 1871, p. 198. 
1867, p. 32. "'The Americaa Matnralisl,' 

■» Ur, ADdei-ion, ' Pmc. Zoiilos. 1873, p- S5. 


354 ^J^ Descent of Man. Pabt U, 

attoek aiem. Noi can I quite disbelieve the several accoimts 
which have appeared of their thus jraialyaing theii prey with fear. 
Some other snakes also make a distinct aoise by rapidly vibinting 
their tails against the surrounding stalks of plants ; and I have my- 
self heard this in the case of a Trigonooephalus in S. America. 

Zoeerh'iio.— The males of some, probably of many kinds of 
lizards fight together from rivalrj. Thns the arboreal AnoK^ 
cristaiellus of S. America is extremely pugnacious : " During the 
"spring and early part of the summer, two adalt males rarely 
" meet without a contest. On first seeing one another, they nod 
" their heads Tip and down three or four times, and at the same 
" time expanding the frill or poach beneath the tbroat ; their 
" eyes glisten with rage, and after waving their tails from 
" side to side for a few seconds, as if to gather enei^, they dart 
" at each other furiously, rolling over and over, and holding 
" firmls with their teeth. The conflict generally ends in one of 
"' the combatants losing his tail, which is often devoured by the 
" victor." The male of this species is considerably larger than 
tlw female;" and this, as fej as Dr. Gunther has been able to 
ascertain, is the general rule with lizards of all kinds. The 
males alone of the CyTiodactyJus rubidas of the Andaman Islands 
possesses pre-anal pores ; and these pores judging from analogy 
probably servo to emit an odour." 

The sexes often di£fer greatly in various esfemal characters. 
The male of the above-mentioned Anolis is furnished with a 
crest which runs along the back and tail, and can be erected at 
pleasure ; but of this crest the female does not exhibit a traea 
In the Indian Cophotis ceylanica, the female has a dorsal crest, 
though much less developed than in the male ; and bo it is, na 
Br. Gunther informs me, with the females of many Iguanas, 
Chameleons, and other lizards. In some species, however, the 
crest is equally developed in both sexes, as in the Iguana tubercu- 
lata. In the genus Sitana, the males alone are furnished with a 
large throat-pouch (fig. 33), which can be folded up like a fan, 
and is coloured blue, black, and red ; but these spl^idid colours 
are eiiibitod only during the pairing-season. The female does 
not possess even a rudiment of this appendage. In the Anolis 
cristateUui, according to Mr. Austen, the throat pouch, which is 
bright red marbled with yellow, is present in the female, though 
in a mdimental condition. Again, in certain other lizards, both 
sexes are equally well provided with throat pouches. Here we 

" Mr, K. L. Austen kept these « Stoliczka, 'Joumsl at Asiatic 

•nimals aliie for a coDsiderable Soc. of BeDgai,' vol. iiiir. 1970, p. 

time; ste 'L:mil ana Water,' July 166, 
1867, p. & 



liepllks u 

o more largely developed i 
a am equally dev loped a both s 
the geaua Draco wh h glida 
thro nb tl Q d r oa their r b- 
Bupported poriohutes and 
which n the boauty of their 
Colo lis haffie description are 
furnifahed w th skinny app n 
da^s to the thioat bke the 
wattl s of gallrna eo 1: rds 
Tl ese become erected when 
tl e ammal 11 exc ted They 
oc r n both sexes I ut a e 
best d veloped wben the male 
arr wt. at matur tj , it wh oh 
age the middle appendage is 
Bometimes twice as long as the head. Most of the species like- 
wise haTe a low crest running along the neck; and this is much 
more developed in the full-grown males, than in the females or 
young males." 

A Chinese species is said to live 
in pairs during the spring ; " and if 
" one is caught, the other falls from 
" the tree to the ground, and allows 
" itself to be captured with impu- 
" nity," — I presume from despair.** 

There are other aud much more 
remarkable differences between the 
sexes of certain lizards. The male 
of Ceratophora aspura bears on the 
extremity of his snout an appendage 
half as loDg as the head. It is 
cylindrical, covered with scales, 
flexible, and apparontiy capable of 
erection ; in the female it is quite 
rudunental. In a second species 
of the same genus a terminal scale 
forms a minute horn on the summit of the flexible appendage ; 

followiag facts in 1 
phora aoi] CbatHicli 
Giinther bimaeif, 

British India,' E 
122, 130, 135. 

" Mr. Swinhc 
Soc.' 1870, p. 24( 

the 'EeptiU 




The Descent of Man 

■pAlST 11. 

and in a third species (C Sloddartn, fig. 3i) the whole appen- 
d^e ia converted into a liorn, which is nsnaDy of a white 
colour, bat assumes a purplish tint when the aEiraal is esdted. 
Tn the adult male of this latter species the horn is half aa inch 
ill length, hut it is of quite minute size in tho female and in the 
vonng. These appendages, as Dr. Giiuther has remained to me, 
may be compared with the combs of galliuaecous birds, and 
sppnrenlly son 

In the genus Chameleon we come to the a«rae of difference 
between the seses. The upper part of the skull of the malo 
€. bi/uTcus (fig. 35), on inhabitant of Madagascar, is produced 
into two great, solid, bony projections, covered with scales Hie 
the rest of the head; and of this wonderful modification of 
Btructuie the fen«Jo exhibits only a rudiment 





Chamcehm Ouienii (fig. 36), from the West Coast of Africa, tlie 
malo bears on his snout and forehead three curious horns, of 
■which the female has not a trace. These horns consist of an 
excrescence of bone coTeied with a smooth sheath, forming part of 
the general int^a- 
ments of the body, 
eo that they are 
identical in struc- 
ture with those of a 
bull, goat, or other 
sheath -homed ru- 
minant. Although 
tho three horns 
differ so much in 
appearance irom 
the two great pro- 
longations of the 
skull m C h/irc s 
wc can hardly doubt 
that they serve tho 
same general pur 
n the economj 

of t 

miJi Theiirstcoji 

jectoro which will occ ir to e cry one is that tl ey are u-^ed 
by the males for fightin^ together and as these animals ire 
very quarrels me " this is probably a correct viei* Mr T ^\ 
Wood also informa me that he once watched two in lividuil of 
(.. pumdrtu, fighting violently on the branch of a tree , they flnng 
their heads about and tried to bite each other ; they then rested 
for a time, and afterwards continued their battle. 

"With many lizards, the sexes differ slightly in colour, the 
tints and stripes of the males being brighter and more distinctly 
defined, than in the females. This, for instance, is the case with 
the above Cophotis and with the Acanthodaetyhta capensis of 
8. Africa. In a Cordylus of the latter country, the male is 
either much redder or greener than the female. In the Indian 
Oaloles niifrihhris there is a still greater difference; the lips also 
of the male are black, whilst those of tho female are green. In 
our common little viviparous hzard (^Zootoco- vivipara) "the 
" under side of the body and base of the tail in the male are 
" bright orange, spotted with black; in the female these parts 
"are pale-greyish-green without spots."™ We have seen that 

'"Dr. Bachdlz, ' Mcnntsbeddit "Bell, 'HisWry of British 

K- Pretas, Atad.* Jan. 1874. p. 78. Reptiles,' 2nd adit. 1849, p. W, 


358 The Descent of Man. PiaTH. 

the niftlee alono of Sitana possess a throat-pouch ; and tJiis ia 
eplendidly tinted with blue, black, and red. In the Proctotrdui 
tenuis of Chile the male alono is marked with spots of blue, 
green, and eoppery-red." In many eases the males retain the 
same colours throughout the year, but in others they liecoiiic 
much brighter during tho breeding-season; I may give as an 
additional instance the C'alofes maria, which at this season has a 
bright red head, the rest of the body being green." 

lloth seses of many species are beaatifnlly coloured exactly 
alike ; and there ia no reason to suppose that such colours are 
protectiye. No doubt with the bright green kinds which live 
in the midst of vegefation, this colour serves to conceal them ; 
and in N. Patagonia I saw a lizard {Frodotrftua multimaadatus) 
which, when frightened, flattened its body, closed its eyes, and 
Uien from its mottled tints was hardly distinguishable from the 
snrrounding sand. But the bright colours with which so many 
lizards ore ornamented, as well as their various curious appen- 
dages, were probably acquired by the males as an attraction, 
and then transmitted either to their male offspring alone, or to 
both sexes. Sexual selection, indeed, seems to have played 
almost as important a part with reptiles as with birds; and the 
less conspicuous colours of the females in comparison with the 
males cannot be accounted for, as Mr. Wallace believes to be the 
ease with birds, by the greater exposure of the females to danger 
during iocubation. 

Second ABT Sexdal Charactbbs of Bmtis. 

Seiual difierenoes — Law of battle — Special weapons — Voctil orgaiu — 

and seasoDnl— Double aod Bingle annual moults— Displiij of oinamenis 
by tlie males. 

Sbconeaby sexual characters are more diversified and con- 
spicuous in birds, though not perhaps entailing more important 
changes of stmctnre, than in any other class of animals. I shall, 
therefore, treat the subject at considerable length. Male bird* 
BomeUmes, though rarely, possess Bpeoial weajKms for lighting 

" For Proctotretns lee ' Zoology the Indian Culotes, nee ' Reptiles of 

of the Voyage of the "Beagie:" British India,' by Dr Giiother, p. 

Keptilei,' by Mr. Bell, p. 8. For 143. 

the Liiards of S. Afric^ lee ' Zoology " Giiather in'Proo. Zoolog. Soc' 

of S. Mica: Beptiles,' by Sir 1S70, p. 778, with « coloured 

Aidrew Smith, pi. 25 and 39. For li|rni«. 


Ghai-. Xin, Birds. 359 

with each, other. They charm the female by TOCal or instni- 
mental music of the most ■varied kinds. They are ornamented 
by all sorts of combs, wattles, protuberances, horns, air-distended 
sacks, top-knots, naked shafts, ptnmea and lengthened feathers 
gracefully springing from all pjurfs of the body. The beak and 
naked akin about tte bead, and the feathers ore often goi^eously 
coloured. The males sometimes pay their court by dancing, or 
by fejitastic antics performed either on the ground or in the air. 
In one instance, at least, the male emits a musky odour, which 
we may suppose serves to charm or excite the female ; for that 
excellent observer, Mr. Bamaay,' says of the Australian musk- 
duck {Biidura Idbata.') that " the smell which the male emits 
" dnrii^ the summer months is confined to that sex, and in 
"some individuals is retained throughout the year; I have 
" never, even in the breeding-season, shot a female which had 
" any smeU of musk," So jmwerful ia this odour durii^ the 
pairing-season, that it can be detected long before the bird can. 
be seen.' On the whole, birds appear to bo the most sestbetic of 
aO animals, excepting of course man, and they have nearly the 
same toate for the beautiful as we have. This is shewn by our 
enjoyment of the singing of birds, and by oni women, both 
civilised and savage, deckii^ their heads with borrowed plumes, 
and usiag gems which are hardly more brilliantly coloured than 
the naked skin aaid wattles of certain birds. In man, however, 
when cultivated, the sense of beauty is manifestly a far more 
comples feeUng, and is associated with various intellectual 

Before treating of the sexual characters with which we are 
here more particularly concerned, I may just allude to certain 
differences between the sexes which apparently depend on 
differences in their habits of life; for such cases, though 
common in the lower, are rare in the higher classes. Two 
hummii^-birds belonging to the genus Enstephanus, which 
inhabit the island of Juan Fernandez, were long thought to bo 
Bpecifically distinct, but are now known, as Mr. Gould informs 
me, to be the male and female of the same species, and they 
differ slightly in the form of the beak. In another genus of 
himimii^-birds (Orypus), the beak of the male is serrated along 
the margin and hooked at the extremity, thus differmg much 
from that of the female. In the Neomorpha of New Zealand, 
there is, as we have seen, a still wider difference in the form of 
the beak in relation to the manner of feeding of the two sexes. 
Something of the same kind has been observed with the gold- 

' 'rbiB,'Toi.iii-;new aeries) 1867, ' Gonld, ' H.-iodbook (0 the Birdj 

p. 4U. of Aostralia,' ISRS, toI. ii. p. 883. 


i6o 77ia Descent of Man. Part n 

finch (Cardmelis e^ejujpB), for I am assured Ijy Mr. J. Jermer'Weii 
that the birdcatohers can distinguish the males by their slightly 
longer beaks. The flocks of males are often found feeding on 
tie seeds of the teazle (Dipsacns), which they can reach with 
their elongated beaks, whilst the ffemales more commonly feed 
on the seeds of the beiony or Serophnlaria. With a slight 
difference of this kiod as a fooudation, we can see how the beaks 
of the two s^es might be made to differ greatly through natural 
Beloction. tn Eome of the above cases, howerer, it is possible 
that the beaks of the males m^ haTe been first modified in 
relation to their contests with other males; and that this 
afterwards led to slightly changed habita of life. 

Law qf BattU. — Almost all male birds are extremely pug- 
nacious, using their beaks, winge, and legs for fighting together. 
We see this ereiy spring with our robins and sparrows. The 
smallest of all birds, namely the humming-bird, is one of the 
most quarrelsome. Mr. Gosse* describes a battle in which a 
pair seized bold of each other's beaks, and whirled round and 
roond, till they almost fell to the ground ; and M. Montea de 
Oca, in speaking of another genus of humming-bird, says that 
two males rarely meet withont a fieree aerial encounter; when 
kept in c^^ "their fighting has mostly ended in the splitting o( 
" the tongue of one of the two, whieh then surely dies from 
"being unable to feed."* With Waders, the males of the 
Eommon wat«r-hen {Qollimda chloropus) "when pairing, fight 
" violently tot the females : they stand nearly upright in the 
" water and strike with their feet." Two were seen to be thus 
ei^ged for half an hour, until one got hold of the head of the 
other, which would haTO been killed, had not the observer 
interfered ; the female all the time looking on as a quiet spec- 
tator.' Mr. Blyth informs mo tliat the males of an allied bird 
(GaUicreoi eristatus) are a third larger than the females, and are 
BO pngnacions during the breeding*season, that thej are kept by 
the natives of Eastern Bengal for the t«ke of fighting. Various 
other birds are kept in India for the same purpose, for instance, 
the bolbnls (PyCTiono(«s hiemorrhous^ which " fight wifh great 

The polygamous ruff (Machi'les pvgnatr, fig 37) is notorious 
for hiB extreme pugnacitj ; and in the spring, the males, which 
ate considerably latter than the females, congregate day after 

Ireland : BiiviB,' tdI. ii. 1850, p 


Chap. Xin. 

Law of Battle. 


day at a particular spot, where the females propose to lay their 
e^s. The fowlers discoyer these spots by the turf being 
tiampled somewhat bare. Here theyflghtTery much like game- 
cocks, seizing each other with thoir beaks and striking with 
their wings. The great ruff of feathers round the neek is then 
erected, and according to Goi. Montagu " sweeps the ground as 
" a shield to defend the more tender parts ;" and this is the only 


362 The Descent of Man. PiUT II 

Instanco known to mo in the case of birds, of any Btructure 
Berring as a shield. The tuff of feathers, howeTer, from its 
varied and rich colours probably serves in chief part as aji orna- 
ment. Like most pt^nacious birds, they eeem always ready to 
fight, and when closely confined often kill each other; but 
Montagu observed that their pugnacity becomes greater during 
the spring, when tbe long feathers on their necks are fuUy 
developed ; and at this period the least movement by any one 
Wrd provokes a general battle.'' Of the pugnacity of web-footed 
birds, two instances will suffice : in Guiana " bloody fights occur 
"during the breeding-season between the males of the wild 
'■ musk-duck (JJairina. twisciuto) ; and where these fights have 
" occurred the river is covered for some distance with feathers." * 
Birds which eeem ill-adapted for fighting ei^age in fierce con- 
flicts ; thus the stronger males of the peUean drive away the 
weaker ones, snapping with their huge beaks and giving heavy 
blows with their wii^. Male snipe fight together, " tugging 
" and pushing each other with their bills in the most curions 
" manner imaginable." Some few birds aro believed never to 
light ; this ia the case, according to Audubon, with one of the 
woodpecfceiB of the United Stat^ {Picus aiiratus), although " the 
" hens are followed by even half a dozen of their gay suitors." ' 

The males of many birds are lai^er than the females, and this 
no doubt is the result of the advantage gained by the larger and 
stronger males over their rivals during many generations. The 
difference in size between the two sexes is carried to an extreme 
point in several Australian species; thus the male musk-duck 
(Bizinra) and the male Cindorttaiphiu cmralis (allied to our 
pipits) are by moasnrement actually twice as large as their 
respective females.'" With many other birds the females are 
laiger than the mal^ ; and as formerly remarked, the explana- 
tion often given, namely, that the females have most of the work 
in feeding their young, will not suffice. In some few cases, as 
we shall hereafter see, the feiaalca apparently have acquired 
their greater size and strength for the sake of conquering other 
females and obtainii^ possession of the males. 

The males of many gallinaceons birds, especially of the poly- 
gamous kinds, are famished with special weapons for fighting 
with their rivals, namely spurs, which ctm be used with fearfal 

' Macgillhray, ' Hiat. BrEt. i. p. 191. For pelicans and eniiifs, 

Birds,' vol. iv, 1852, pp. 177-181. se* vol. iii. pp. 138, 477. 

' Sir E. Solioinbnrgk, in ' Jourusl " Geulil, ' Handbook of Birds 01 

ofR.Geograpli.Soo.'vol.iili. 1843, Australia,' vol. i. p. 3ffi ; toI ii. p. 


CaiP. XIII. Law of Battle. 363 

effect. It h^ been recorded by a trustworthy writer " that in 
Derbyshire a Mte Btrnck at a. game-hen accompanied by her 
chickens, when the cock rushed to the rescue, and drOTO his 
epur right throngli the eye and ekull of the a^reesor. The 
Bpur wBB with difficult? drawn from the skull, and as the kite 
though dead retained his grasp, the two birds were firmly 
locked together; but the cock when disentangled was very 
little injured. The invincible courage of the game-cock ia 
aotorioUB : a gentleman who loi^ ago witnessed the brutal 
scene, fold me that a bird had both its legs broken, by some 
accident in the cockpit, and the owner laid a wager that if the 
l^js could be spliced so that the bird could stand upright, he 
would continue fighting. This was effected on the spot, and the 
bird fought with undaunted courage until lie received his death- 
stroke. In Ceylon a closely allied, wild species, the Oallas 
Blanlejfi, ia known to flglit desperately "in defence of his 
" seraglio," so that one of the combatants is frequently found 
dead.'" An Indian partridge {Vrtygamie ffuione), the male of 
which is furnished with stroi^ and sharp spurs, is so quarrel- 
some, " that the sears of former fights disfigure the breast of 
" almost every bird yon Idll," ^ 

The males of almost all gallinaceous birds, even those which 
are not ttimished with spurs, engage during the hreedii^-season 
in fierce conflicts. The Capercailzie and Blaek-cock (Teirao 
urogallua and T. tetrix), which are both polygamists, have regular 
appointed places, where during many weeks they congregate in 
numbers to fight tt^ther and to display their charms before the 
females. Dr. W. KovaJevskj informs me that in Russia he has 
BOin the snow all bloody on the arenas where the capercailzie 
haTe fought ; and the black-cocks " make the feathers ily in every 
" direction," when several " engage in a battle royaJ." The 
elder Brehm gives a curious accoimt of the Balz, as the love- 
dances and love-songs of the Black-cock ace called in Germany. 
The bird utters almost continuously the strangest noises; " he 
" holds his tail up and spreads it out like a fan,"he lifts up his 
" head and neck with all the feathers erect, and stretches his 
" wings from the body. Then he takes a few jumps in different 
" directions, sometimes in a circle, and pressos the imder part of 
" his beak so hard against the ground that the chin feathers are 
" rubbed ofE During these movementE he beats his wings and 
" turns round and round. The mora ardent he grows the more 
" lively he becomes, untQ at last the bird appears like a frantio 

" Mr. H*witt in the 'Poultry Nat. Hist.' vol. liy. 1854, p. 63. 
Bnok by Teeetmeier,' 1866, p. ]37. " Jerdon, 'Birils of India..' tcL 

'' Iiaysrcf, 'AnnaU and Mag. of iii, p. 5''4. ■ 


j64 "^he Descent of Man. Pabt TL 

" creatnie." At snch tiroes the Wack-cocks aro so absorbed that 
they become almost blind and deaf, but less so than the caper- 
cailzie : hence bird after bird may be shot on the same spot, or 
even caught by the hand. After performing these antics the 
males b^in to ^ht : and the same black-eocfc, in order to prove 
hia strength over several antagonists, will yistt in the course of 
one morning several Balz-places, which remain the same during 

The peacock with his long train appears more like a dandy 
than a warrior, but he sometimes eng^es in fierce contests : the 
Bev. W. Darwin Toi informs me that at some tittle distance 
from Chesifir two peacocks became so excited whilst fighting, 
that they flew over the whole city, still engaged, until they 
alighted on the tflp of St. John's tower. 

The spur, in those gallinaceous birds which are thus provided, 
is generally single ; but Polyplectron (see fig. 51, p. 397) has two 
or more on each 1^; and one of the Blood-pheasants (Ithaginis 
cruemfos) has been seen with five spurs. The spurs ai-e generally 
confined to the male, being represented by mere kuobs or rudi- 
ments in the female ; but the females of the Java peacock (Pavo 
miilkus) and, as I am informed by Mr. Blyth, of the small fire- 
backed pheasant (Eiiplocamiis erytkropthalmus) possess spurs. 
In Galloperdii it is usual for the males to have two spurs, and 
for the females to have only one on each leg.'° Hence spura may 
be considered as a masculine structure, which h£ffi been occasion- 
ally more or less transferred to the females. Like most other 
secondary sexual characters, the spurs are highly variable, both 
in number and development, in the same species. 

Various birds hare spurs on their wings. But the Egyptian 
goose (Chenalnpe^: legypHaeiis) has only " bare obtuse knobs," and 
these probably shew us the first steps by which true spurs have 
been developed in other species. In the spur-winged goose, 
Flectrnjderus yambensis, the males have much larger spurs than 
the females; and they use them, as I am informed by Mr. 
Bartlett, in ^Jiting bother, so that, in this case, the wing-spurs 
serve as sexual weapons ; but according to Livingstone, they are 
chiefly used in the defeUM of the yonng. The Palamedea 
(fig. S8) is armed with a pair of spurs on each wii^ ; and theKo 
are such formidable weapons, that a single blow has been known 
to drive a dog howling away. But it does not appear Ibat the 
spurs in this case, or in that of some of the spur-winged rails, 

" Brehr 

n, 'Illust. Thierleben, 

Sweden,' &c, 1867, p. 79. 

1867, B. 1. 

-. fL 351. Some of the 

" Jerdon, ' Bltis of India; on 



Ith/LStinis, ™l. Hi, p. S33; cd Gall* 


LalJO of BattU. 


are larg in tj.e mal than m the female ^ In ceriain plovers^ 
however the wiiiij spura must be coHBiiiered as a eexual oba- 


3(56 The Descmt of Man. Part 11 

meter. Thus in the male of our cammon peewit ( Vomdlvx cW9- 
tatus) the tubercle on the shoulder of the wing becomes more 
prominent during the breedit^-season, and the males fight 
blether. In some species of lobiyanellus a similar tubercle 
becomes developed during the breeding-season "into a short 
" homy spur." In the Australiaoi. ^iaius both sexes haYeBpurs, 
but these are much larger in the males than in the females. Iii 
an allied bird, the Hoplopterm a/rmat is the spurs do not increase 
in size durii^ the breeding-season but these birds have been 
seen in Egjpt to fight together m the san o manner as our 
peewits, bj turning suddenly in the air ani t.trikmg siievays 
at each other, sometimes with fatal lesulta Ihus xIbo they 
drive away other enemies," 

The season of love is that of 1 xttln but the n ales of sjmo 
birds, as of the game-fowl and ruff and even the yonog males of 
tlie wiid turkey and grouse," are leody to tight whenever they 
meet. The presence of the female is the te f m/ bell circii 
The Bengali baboos mate the pretty little males ol thp ama lavit 
(Estrtlda amnnditva') fight t(^thor by placing three small cages 
in a row, with a female in the middle; after a little time the two 
males ajre turned loose, and immediately a desperate battle en- 
sues." When many males congregate at the same appointed 
spot and fight together, as in the case of grouse and various other 
birda, they are generally attended by the females," which after- 
wards pair with the victorious combatants. But in somecases tho 
pairing precedes instead of succeeding the combat : thus accord- 
ing to Audubon," several males of the Virginian goat-sucker 
fS^aprimulgas virginian'm) " court, in a highly entertainin:; 
" manner the female, and no sooner has she made her choice, 
" than her approved gives chase to all intmders, and drives 

meJea, Brehm's 'Tliisriebea,' B. jv. 

" Kithnnls^D on Tetrtio HmielMt 

t. T40. See nlao on thi? birii Azitra, 

'I'iiuna Bor. Amer.: ilirJV 18:fl, 

'Voj-iigfls daua rAmerique meild.' 

p. 343. L Lloyd, 'Game Birds o( 

to™, iv. 180a, pp. 179, 253. 

Swedsa,' 1867, pp. 22, 79, on the 

" See, oa our peewit, Mr, R. Can 

capercailzie and black-cock. Brehm, 

m 'Land and Water,' Aug. Stti,, assert* (' Thiwieben,' io.. 

1368, p. 48. In regard to Lobi- 

B. it. a. 352) thai in (^etmanv the 

vanellna, see Jerdon'a 'Biriii of 

grej-heos do not gentrally attend 

ladia,' vol. iii p. 647, find Gould's 

the Balzen of the black-coch^ but 

this <s an eiceptioa to the commoj 

Tol. ii. p. 220. For ttte Holoptems, 

rule; possiblj the hcna may lie 

jM Mr. Allen ia the 'Ibis,' toI. t, 

hidden in the surrounding bu>ihes. 

lSii3, p. 156. 

as IS knonn to be the case with the 

" Audub™, ' Oruith. Bii^raphr, 

grey-hens in Scandinavia, and with 

vol. ii. p. 492; vol. 1. pp. 4-13. 

'• Mr. Blyth, Land and Water, 

I8G7, p. 213. 



Chap. SHI. Law of Battle. 36; 

" them beyond his dominions." Generally tlie males try to drira 
awiif 01 kill their rivals before they pair. It does not, however, 
appear that the females invariably prefer the victorions males. 
I have indeed been, assured by Dr. W. Kovalevsfcy that the 
female capercailzia sometimes steals away with a young male 
who has not dared to enter the arena with the older codes, in 
the same manner as occasionally happens with the does of the 
red-deer in Scotland. When two mal^ contend in presence ot 
a sii^le female, the victor, no doubt, commonly gains his desire ; 
but some of these battles are caused by wandering males trying 
to distract the peace of an already mated pair.™ 

Even with the most pagnacious species it is probable that the 
pairing does not depend esclusively on the mere strength and 
courage of the male ; for such males are generally decorated with 
various ornaments, which often become more brilliant during the 
breeding-season, and which are sedulously displayed before the 
females. The males also endeavour to charm or excite their 
mates by love-notes, songs, aud antics ; and the courtship is, in 
many instances, a prolonged affair. Hence it is not probable 
that the females are indifferent to the ohanns of the opposite 
sei, or that they are invariably compelled to yield to tho victorious 
males. It is more probable that the females are excited, either 
before or after the conflict, by certain males, and thus im- 
consciously prefer them. In the case of Tstrao umbeUiis, a good 
observer" goes so far as to believe that the battles of the males 
" are all a sham, performed to show themselves to tho greatest 
" advantage before the admiring females who assemble around ; for 
" I have never been able to find a maimed hero, and seldom more 
" than a broken feather," I shali have to recur to this subject, 
but I may here add that with the Tetrim eupido of the Uniteo 
States, about a score of males assemble at a particular spot, ana 
strutting about, make the whole air resotmd with tlieir extra 
ordinary noisee. At the first answer from a female the males, 
begin to fight furiously, and the weaker give way ; but then, 
accordii^ to Audubon, both the victors and vanquished search 
for the female, so that the females must either